ORNITOLOGIA NEOTROPICAL 13: 381–396, 2002© The Neotropical Ornithological Society

STATUS AND CONSERVATION OF THE AVIFAUNA OF THE YAXCHILÁN NATURAL MONUMENT, CHIAPAS, MÉXICO

Fernando Puebla-Olivares, Emir Rodríguez-Ayala, Blanca E. Hernández-Baños, & Adolfo G. Navarro S.

Museo de Zoología, Facultad de Ciencias, Universidad Nacional Autónoma de México, Apartado Postal 70-399, México, D.F. 04510, México.

Resumen. – Estatus y conservación de la avifauna del Monumento Natural Yaxchilán, Chiapas,México. – Se presenta una lista de la avifauna encontrada en el área protegida de Yaxchilán, Chiapas,México. Un total de 235 especies fue registrado en la zona, 75.7% de las cuales fueron residentes perma-nentes, 19.6% residentes de invierno, y 4.8% residentes de verano o pertenecientes a otras categorías esta-cionales. Dado el buen estado de conservación de la zona, se registran poblaciones importantes deespecies catalogadas en alguna categoría de amenaza como el Águila Arpía (Harpia harpyja) y otras, de lascuales se hace mención. Se mencionan además registros importantes para la avifauna mexicana como unejemplar mexicano del Fandanguero Pechiescamoso (Phaeochroa cuvierii) y el primer registro en México delHormiguero Calvo (Gymnocichla nudiceps).

Abstract. – An avifaunal list of the Yaxchilán Natural Monument, Chiapas, Mexico, is presented. A totalof 235 species was recorded, 75.7% of which were permanent residents, 19.6% winter residents, and 4.8%summer residents and of other seasonal categories. Given the pristine status of the area, populations ofglobally endangered species such as the Harpy Eagle (Harpia harpyja), among others, were recorded and arementioned. The area provided new and interesting records for the Mexican avifauna, including a specimenof Scaly-breasted Hummingbird (Phaeochroa cuvierii) and the first Mexican record of Bare-crowned Antbird(Gymnocichla nudiceps). Accepted 22 February 2002.

Key words: Avifauna, Yaxchilán, Chiapas, Mexico.

INTRODUCTION

Yaxchilán, a protected area located in theState of Chiapas, México, has a great value forconservation. First because it is located in theeasternmost section of the state of Chiapas, inone of the most important remains of pristinetropical vegetation of the country. Alsobecause, as in several areas of northern Cen-tral America, some of the sites where conser-vation of biological systems has been possibleare also very valuable for containing impor-tant archaeological Maya sites. Thus, efforts

are made to preserve their biological and cul-tural values entirely. Yaxchilán is located in theCentral Maya Lowlands, an area where Mayaculture arose in the Preclassic Period, duringthe Olmec Empire splendor in eastern Mex-ico, and whose cultural influence extendedsouth to Costa Rica (SEMARNAP-INE-CONABIO 1995).

The biological diversity found in this rela-tivelly small area is astonishing, in spite ofbeing relatively dryer than the adjacent rainforests of Chajul and Montes Azules(González-García 1993). Therefore, efforts

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should be directed to make a detailed study ofthe biological richness of the region, and pro-mote its conservation and educational value.

The results presented herein are part of amajor collaborative research among biologistsand anthropologists to evaluate biologicaldiversity of the area, and establish the basisfor a more general conservation plan. Thus,the main objective of this contribution is toprovide a list of the birds of the region,highlighting noteworthy species and adding

information for future conservation strate-gies.

STUDY AREA AND METHODS

The Yaxchilán Natural Monument is locatedat a bend of the Usumacinta River (16o53’N,90o58.60’W; Fig. 1). The official boundariesare specified in a Presidencial Ordinance of24 August 1992 that preserved an area of2621 hectares, most of it “ejidos” and com-

FIG. 1. Geographic location of the “Monumento Natural Yaxchilán” in Chiapas, Mexico.

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munal land, although the archaeological zoneis federal property (SEMARNAP-INE-CONABIO 1995). The archaeological zoneis now under care of the National Institute ofAnthropology and History (INAH) and hasadequate installations for housing personnel.Access to the zone is by air, or by land arriv-ing to Palenque, and from there to the townFrontera Corozal, then by boat through theUsumacinta River to Yaxchilán.

Yaxchilán is part of the Lacandona Forest,one of the most inaccessible and biologicallyunknown regions from Chiapas, that hasbeen assigned to the “Eastern Highlands”phytogeographic region of Breedlove (1970),where several low montane zones declinegradually toward the drainage of the Usuma-cinta River. The soils of the Lacandonaregion are constituted of limestones, withgritty and volcanic extrusions, and with arange of elevation from 400–1500 m. Vegeta-tion is relatively uniform, with tropical rainforest being the most common type. How-ever, patches of low savanna, as well as dis-persed palm-tree forests, are common. Therain forest is present in the flat areas at theupper drainage of the Usumacinta. The flo-ristic associations of this area are continuouswith those of the Petén in Guatemala, bothcontaining a large number of endemic plantspecies (Breedlove 1970).

At Yaxchilán, the dominant vegetationtype is tropical rain forest (Meave del Castilloet al. in prep.), but fragments of secondaryvegetation are common in areas adjacent tothe archaeological zone and the camp of theINAH. The vegetation that is found alongthe course of the Usumacinta River is knownin the zone as “jimbal,” because of the pres-ence of Guadua sp., and is characterized bythe presence of plants similar to “carrizo”.Three arboreal strata are present. The highestis composed of trees 50–60 m in height withstraight and butressed trunks emergent abovethe canopy. A medium-height stratum forms

a continuous canopy to a height of 25–40 m.The third stratum is composed of trees withbranches to a height of 10–20 m. Understoryshrubs and grasses are practically absent andthe epiphytes are present only in the higheststratum (Breedlove 1970, Pennington &Sarukhán 1998). Some of the most commontrees in the canopy are Apidosperma megalo-carpa, Brosimum alicastrum, Dialium guianense,Swietenia macrophylla, Erblichia xilocarpa, Guatte-ria anomalous, Manilkara achras, Poulsenia armataand Terminalia amazonia. The common treesof the subcanopy are Alchornea latifolia, Aliber-tia edulis, Belotia cambelli, Bumelia persimili, Cassiagrandis, Blepharidium mexicanum, Bursera sima-ruba, Guarea excelsa, Hasseltia dioica, Licariapekii, Orthion subsessile, Pithecelobium arboreum,Quararibea funebris, Wimmeria bartletii andZuelania guidonia (Breedlove 1970).

Field work was performed by two to fiveresearchers every two months, from Decem-ber of 1997 to February of 1999 completing atotal of 70 days of fieldwork. Surveys of theavifauna were made using mist-nets (approx.8040 mist net hours) and gathering observa-tional and sound records from the area.Selected specimens were obtained of as manyspecies as possible to construct a vouchercollection of skins, skeletons, and frozen tis-sue samples, deposited at the Museo deZoología “Alfonso L. Herrera”, Facultad deCiencias, Univ. Nacional Autónoma de Méx-ico (MZFC), in Mexico City. Data for speci-mens and sight records were included in adatabase elaborated by using the BIOTICA2.0 software of CONABIO (1997) for easymanagement.

An estimate of the abundance of eachspecies during the study was obtainedthrough a classification of the percentage ofthe total field days (70 for residents, 42 formigrants) in which a species was recorded.Four classes are used: Rare (1–7%), uncom-mon (8–29%), common (30–57%), and verycommon (58–100%); relative abundance

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assignation for each species is depicted inAppendix 1.

Additional information on the region'savifauna was obtained from Howell (1989)and the few previously collected specimensfrom the area. Those specimens are housed inthe Canadian Museum of Nature, Ottawa,Canada (CMN, six specimens) and theNational Natural History Museum in Paris,France (MNHNP, 209 specimens), all col-lected by the J. L. Mottron and R. Magrisexpeditions between 1974 and 1976 (C. Joua-nin pers. com.). Taxonomy follows AOU(1998).

RESULTS

Yaxchilán is a site of high bird diversity. Ofthe approximately 1060 species of birdsrecorded in Mexico, 425 (40.1%) have beenrecorded in Chiapas, and 222 (20.94%) wererecorded in the locality of Yaxchilán by us,plus thirteen extra species recorded only by

specimens in other museums, e.g., the Black-billed Cuckoo (Coccyzus erythrophthalmus), theFerruginous Pigmy Owl (Glaucidium brasi-lianum), and the Lincoln Sparrow (Melospizalincolnii) in MNHNP (Appendix 1), for a totalof 235 species. Thus, of the total of speciesrecorded in Chiapas, 55% occur in the rela-tively small area of Yaxchilán (2621 ha) incomparison with larger nearby reserves e.g.,Montes Azules (300 species in 331,200 ha),Calakmul (235 species in 723,185 ha), ElOcote (more than 350 species in 48,140 ha),or Monumento Natural Bonampak (300 spe-cies in 4357ha) (SEMARNAP-INE-CONA-BIO 1995).

Analysis of the accumulation curve forspecies recorded by dates of field work wasadjusted through an exponential growthmodel employed to compare predictednumber of species (Gutiérrez 1984). This esti-mate suggests that most of the avifauna thatoccurs at the site was recorded (Fig. 2,Appendix 1).

FIG. 2. Accumulation curve of species recorded and hypothetical total number of species (Gutiérrez1984) in Yaxchilán.

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Species recorded belong to 45 families of17 orders of birds, 75.7% of which (178 spe-cies) are permanent residents, 19.6% (46 spe-cies) are winter residents, and 4.8% are eithersummer residents or accidental in the region(11 species). The Tyrannidae family showsthe highest species richness (27 spp.),followed by Parulidae (20 spp.), Thraupidae(15 spp.), Icteridae (12 spp), Trochilidae (11spp), Cardinalidae (10 spp.), Accipitridae (9spp.), Ardeidae (8 spp.), Picidae (7 spp.), Den-drocolaptidae (7 spp.) and Columbidae (7spp.). A voucher collection of 531 specimensfrom 108 species was constructed (Appendix1).

The distribution of species richness byseasonal status through the year showed aninteresting pattern. The highest richness ofpermanent and winter residents was recordedin February and April, while lowest valueswere obtained in June (Fig. 3).

Added value to the conservation of birdsof the area is the presence of an important setof species that are classified as threatened

either globally, [e.g., Harpy Eagle (Harpia har-pyja) and Orange-breasted Falcon (Falco deiro-leucus) (Collar et al. 1992, BirdLifeInternational 2000)], or nationally as threat-ened or endangered (21 species, DOF 1994,Appendix 1).

Given the pristine status of conservationof vegetation at Yaxchilán, and the relativelypoor ornithological knowledge of the area,some species are noteworthy and are presentsometimes with important and healthy popu-lations.

SPECIES ACCOUNTS

King Vulture (Sarcoramphus papa). Thisspecies is a rare resident of the area and wasonly recorded on 10 and 11 February 1998in a patch of grass adjacent to the ruins,and on 20 August 1998 at the UsumacintaRiver.

Harpy Eagle (Harpia harpyja). On 6 and 7April, and 26 1998 June, we observed one

FIG. 3. Species richness by seasonal status in the different field trips.

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individual of this highly endangered eaglesoaring low over the canopy of well-preservedrain forest. This species is typically associatedwith pristine vegetation, and its presence indi-cates the good situation of the habitat. This isone of the scarce recent records of the speciesin Mexico (Escalante & Peterson 1993,Morales-Pérez 1998) and it is considered rarein the area.

Black Hawk Eagle (Spizaetus tyrannus). Thisspecies is a rare resident in the area and wasobserved on 10 February and 23 and 25August 1998, perching at the top of a tall treeat the rain forest edge close to the ruins.

Ornate Hawk Eagle (Spizaetus ornatus). Thiseagle is also a rare resident in the region, onlyone individual was observed perching insidethe canopy of well-preserved rain forest on 31October 1998.

Orange-breasted Falcon (Falco deiroleucus). Thisfalcon is considered an uncommon residentof tropical rain forests in Middle America.However, it is a rare resident in the localitybecause only one individual was observedperching in a dead tree close to the ruins on 7February 1998. This record is the first pub-lished for Chiapas and one of few records forMexico. Although Howell & Webb (1995)mention that the only known specimens fromMexico come from Tecolutla, Veracruz,other specimens are known from Chiapas(Ocosingo, FMNH).

Crested Guan (Penelope purpurascens).Healthy populations of this declining cracidwere observed during several months of theyear, flying and foraging at the medium-height stratum of the forest. In the area it isan uncommon species; however, local peopleinformed us that it is more commonlyobserved during the dry season (october tomay).

Great Curassow (Crax rubra). Although notrecorded directly by us, locals informed usthat scattered individuals are present at Yax-chilán during the dry season. The species wasrecorded from nearby areas in Montes Azulesby González-García (1993).

Scarlet Macaw (Ara macao). Groups of 30 to 40individuals were observed year-round flyinghigh up the canopy. We considered this spe-cies common in the area; however, given theillegal trade in this macaw, Yaxchilán andnearby reserves in northern Chiapas andGuatemala might represent one the few ref-uges for this species in northern CentralAmerica (Iñigo-Elías 2000).

Short-tailed Nighthawk (Lurocalis semitorqua-tus). This rare and elusive nightjar was firstsight recorded for Mexico by Howell (1989)at Yaxchilán, with further observations inother sites in Chiapas and in northern CentralAmerica (no specimens available yet, Howell& Webb 1995). No records of this specieswere obtained by us during the field work,although a special effort was devoted tolocate it in suitable habitat. The seasonal andtaxonomic status of the populations in Mex-ico, Guatemala, and Honduras remainunknown.

Scaly-breasted Hummingbird (Phaeochroa cuvi-erii). First recorded in Mexico by Feltner(1976) and Howell (1989) who recorded thespecies in breeding condition at Yaxchilán,and later by González-García (1993) inthe Montes Azules Biosphere Reserve. Onemale (MZFC 14887) was collected on 10 Feb-ruary 1998, in a patch of rain-forest, consti-tuting one of the three known Mexicanspecimens. The other two were collected by J.L. Mottron (Allan R. Phillips' collection) atYaxchilán on 22 October 1974 and are depos-ited at the Canadian Museum of Nature,Ottawa.

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Buff-bellied Hummingbird (Amazilia yucatanen-sis). One specimen (MZFC 14888) was col-lected on 3 November 1998. This record isthe southernmost for the species in Mexicoand expands its known distributional areasouth to eastern Chiapas.

Bare-crowned Antbird (Gymnocichla nudiceps).This antbird inhabits the tropical rain forestsof eastern Guatemala and southern Belize toHonduras. An adult male of this species wasobserved and unmistakably identified atYaxchilán in November 1996, foraging inthe undergrowth of dense primary rainforest. This is the first Mexican record ofthe species and represent a range extensionof approximately 223 km northwest ofits closest known locality at the Belize-Guatemala border [based on Howell &Webb's (1995) map for the species].The species was not observed again in sub-sequent visits, which suggests that it is veryrare or that its presence in Mexico is acciden-tal.

Northern Royal Flycatcher (Onychorhynchus“coronatus” mexicanus). This species is oftenconsidered as an uncommon resident ofhumid lowland forests, especially associatedwith rivers (Howell & Webb 1995). The spe-cies is an uncommon resident in the area,inhabiting the dense primary vegetation, for-est edges, trails, and semiopen areas near theruins. Specimens were obtained year round(MZFC 14725-29, Feb to Sept 1998). Thepopulation in Yaxchilán might constitute oneof the most important in northern CentralAmerica because the specie is typically associ-ated with pristine vegetation.

Lovely Cotinga (Cotinga amabilis). Individualsof this uncommon species were observedforaging on fruits of a Ficus tree near theruins several times in December 1997 andAugust 1998.

White-browed Wren (Thryothorus albinucha).One specimen (MZFC 14633) was collectedon 24 August 1998, partly filling the distribu-tional gap depicted in Howell & Webb (1995)between northeasternmost Chiapas and Gua-temala.

Green Honeycreeper (Chlorophanes spiza). Fourspecimens of this uncommon species werecollected on 7–9 April 1998 (MZFC 14427-28) and 14–15 December 1997 (MZFC144429-30). The individuals were foragingon fruits of a vine in a Ficus tree aspart of a mixed-species flock in the loweststratum of the tree canopy near the ruins.Additional specimens were collected in 1974(MNHNP).

Blue Seedeater (Amaurospiza concolor). This isan uncommon and very local resident of themid-elevation brushland and forests, oftenassociated with bamboo (Howell & Webb1995). Records of this species in MiddleAmerica are very scarce and seldom pub-lished. One adult male of this rare and elusiveseedeater was obtained in a patch of jimbal(Guadua sp.) on 29 August 1998 (MZFC14736). This record expands its known distri-bution 160 km northeast from the closestknown locality in Chiapas (Socoltenango,MLZ 35253).

Blue Bunting (Cyanocompsa parellina). Onespecimen (MZFC 14434) was obtained on 23June 1998, expanding the known range of thespecies in Mexico approximately 63 km southfrom the outlined distribution of Howell &Webb (1995) in eastern Tabasco.

Spot-breasted Oriole (Icterus pectoralis). Threespecimens collected in 1974 by the Mottron-Magris expedition (MNHNP 1975.974-76)constitutes de northernmost records of thisspecies in Chiapas, mainly restricted to thePacific lowlands of Middle America and occa-

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sional in the central valleys of Guatemala(Howell & Webb 1995).

DISCUSSION

The high species richness found at Yaxchilánis likely to increase as more survey effort isdevoted. Although predictive models suggestthat most of the avifauna has been recorded,special effort should be devoted looking fornocturnal birds (e.g. Bubo, Chordeiles), transientmigrants [e.g., Swainson's Hawk (Buteo swainso-nii), Broad-winged Hawk (B. platypterus), East-ern Kingbird (Tyrannus tyrannus), ChimneySwift (Chaetura pelagica), Red-eyed Vireo (Vireoolivaceus), and Gray-cheeked Thrush (Catharusminimus)], and other winter residents thatmust be present in the region but have beenunrecorded for lack of surveying in the rightseason of the year. With the informationavailable, however, it is possible to discuss theimportance of the region for knowledge ofMexican birds and for conservation.

The avifauna of Yaxchilán is unique inseveral ways, especially because geographiclocation and avifaunal data suggest that it mayrepresent the contac zone of different sets ofbird faunas. First, it is characterized by smallpopulations of species found nowhere else inMexico [e.g., Short-tailed Nighthawk (Luroca-lis semitorquatus), White-whiskered Puffbird(Malacoptila panamensis), Bare-crowned Ant-bird, Scaly-breasted Hummingbird], thatbelong to the Central American Atlanticmoist forest ecoregion (Dinerstein et al. 1995)and might reach the easternmost Chiapas intheir western distributional limit. Other spe-cies with similar distribution patterns [e.g.,Crested Eagle (Morphnus guianensis), Oliva-ceous Piculet (Picumnus olivaceus), White-winged Becard (Pachyramphus polychopterus)],“hearsay” recorded in Mexico for long time,may well be present in the region as casuals.Finally, several species characteristic of thedryer semideciduous tropical forest of the

Yucatan Peninsula (e.g., White-browed Wren,Buff-bellied Hummingbird) found here theirsouthern distributional limit. Those speciescoexist widely with the species's rich avifaunaof the rain forest of the Isthmus of Tehuante-pec, and the Pacific lowlands (e.g.,Spot-breasted Oriole). Thus, from the biogeo-graphic point of view, Yaxchilán and nearbyareas are a region that deserves deeper studiesinvolving full biotas for explaining the causesof such a complex species composition.

Yaxchilán is also unique in having healthypopulations of rain-forest species considereduncommon or rare in other regions of thecountry [e.g., Grey-chested Dove (Leptotilacassinii), White-whiskered Puffbird, NorthernRoyal Flycatcher, Rufous-tailed Jacamar (Gal-bula ruficauda). At the same time, some healthypopulations of species with a conservationstatus are also present [e.g., Scarlet Macaw,Harpy Eagle, Solitary Eagle (Harpyhaliaetussolitarius)]; see Species Accounts). It is also ofinterest to note the almost complete absenceof human-related species like the House Spar-row (Passer domesticus), or cowbirds (Molothrusspp.), and the rarity of Great-tailed Grackles(Quiscalus mexicanus). These data supplyimportant information about the good con-servation status of the habitats of the region,and their importance as a key conservationarea for a very particular fauna in northernCentral America.

The complexity in composition, structureand function of the tropical rain forests, makethem very vulnerable to human perturbation.In the last 30 years, the original area of thetropical rain forest in Mexico has been drasti-cally reduced, with less than 10% left in goodpreservation condition (Flores-Villela &Gerez 1994). Habitat destruction is advancingat a very fast rate, human settlements aremore numerous, and many taxa are exploitedfor the illegal pet trade (e.g., many of the par-rot taxa found in the area). To be successful,conservation efforts in the tropics should

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become a multidisciplinary issue, involvingtaxonomists, ecologists, anthropologists,economists, and politicians (SEMARNAP-INE-CONABIO 1995) working with localpeople to develop action plans and strategiesfor preservation and use of biological diver-sity.

Biological information at several levels ofbiodiversity studies (taxonomic, ecological,and genetic) is strongly needed for amultitude of protected areas in the world.While many efforts are devoted recently todetect, with biological criteria, priority areasfor bird conservation in the world (e.g., Wege& Long 1995) and in Mexico (e.g., Arizmendi& Márquez 2000), some areas have beendeclared as important for conservation,although they lack a biological survey thataccounts for its biological importance.

Although we presented only the resultsobtained in the bird survey, similar informa-tion exists for mammals, reptiles, amphibians,insects, and flora, that will be published else-where (Meave & Luis in prep.). This model ofgathering information from different sources,and analyzing it in a broad context, has beensuccessful in Mexico for several preservedareas, e.g., Omiltemi, Guerrero (Luna &Llorente 1993), Los Tuxtlas, Veracruz(González-Soriano et al. 1997), and the Sierrade la Laguna and El Vizcaíno, Baja CaliforniaSur (Arriaga & Ortega 1998, Ortega &Arriaga 1991). However, the particular situa-tion of Yaxchilán, on the border of Mexicoand Guatemala, should enhance the develop-ment of inventory and conservation efforts ata multi-national scale, improving the knowl-edge and conservation of natural regionsacross borders.

ACKNOWLEDGMENTS

Jorge Meave del Castillo and Armando Luis-Martínez coordinated the group activitiesduring the project. We also thank Livia León,

Armando Luis, Samuel López, Sergio Larios,Luis Antonio Sánchez, and Ubaldo Melo forinvaluable assistance in the field. We thankIsolda Luna, Fanny Rebón, RaymondMcNeil, Guy Kirwan, and two anonymousreviewers, for comments on the manuscript.We also thank the late Henri Ouellet andMichel Gosselin (Canadian Museum ofNature, Ottawa;CMN); Christian Erard,Christian Jouanin, and Francis Roux(Museum National d'Histoire Naturelle,Paris); John Hafner (Moore Laboratoryof Zoology, MLZ); Mercedes Foster (UnitedStates National Museum,USNM); A.Townsend Peterson, John Bates, and DavidWillard (Field Museum of Natural History,FMNH), for access to specimens intheir care. Claudia Abad, Alejandro Gordillo,Hesiquio Benítez, Elsa Figueroa, andAlad Flores helped in obtaining crucialspecimen information in Paris and Ottawa.Financial support was obtained from theComisión Nacional para el Conocimiento yUso de la Biodiversidad (CONABIO grantNo. M099), CONACyT, DGAPA- UNAM,and National Science Foundation. Specialthanks to the Instituto Nacional deAntropología e Historia (INAH), thatkindly supplied enormous logisticsupport and access to their ResearchStation. The Instituto Nacional de Ecología(INE) provided the collecting permits.

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Ortega, A., & L. Arriaga. 1991. La reserva de laBiósfera El Vizcaíno en la Península de BajaCalifornia. Publ. 4, Centro de InvestigacionesBiológicas de Baja California Sur, México, D.F.

Pennington, T. D., & J. Sarukhán. 1998. Árbolestropicales de México. 2nd ed. Univ. NacionalAutónoma de México y Fondo de CulturaEconómica, México, D.F.

SEMARNAP-INE-CONABIO (Secretaría deMarina Recursos Naturales y Pesca-InstitutoNacional de Ecología-Comisión para el Cono-cimiento y Uso de la Biodiversidad). 1995.Reservas de la biósfera y otras áreas naturalesprotegidas de México. Secretaría del MedioAmbiente Recursos Naturales y Pesca, InstitutoNacional de Ecología y Comisión Nacionalpara el Conocimiento y Uso de la Biodiver-sidad, México, D.F.

Wege, D. C., & A. Long.1995. Key areas for threat-

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ened birds in the Neotropics. Birdlife Conser-vation Series No. 5, BirdLife International,

Cambridge, UK.

APPENDIX 1. Systematic list of the birds of Yaxchilán. Seasonal status (SS) codes are: WR, winter resi-dent; T, transient; O, occasional; SR, summer resident; PR, permanent resident. Conservation status (CS)codes according to the Mexican Endangered and Threatened Species List (DOF 1994) are: A, threatened;R, rare; PE, endangered; SPE, subject of special protection. Records: O (observed), C (collected anddeposited at MZFC), M (other museum specimens). Abundance: R (rare), U (uncommon), C (common), V(very common).

Common names Species SS CS Records AbundanceGreat Tinamou Little TinamouThicket TinamouSlaty-breasted TinamouBrown PelicanNeotropic CormorantGreat Blue HeronGreat EgretSnowy EgretLittle Blue HeronTricolored HeronCattle EgretGreen HeronBlack-crowned Night-HeronBlack VultureTurkey VultureKing VultureBlack-bellied Whistling DuckBlue-winged TealOspreyPlumbeous KiteWhite HawkCommon Black HawkGreat Black HawkSolitary EagleRoadside HawkHarpy EagleBlack Hawk EagleOrnate Hawk EagleBarred Forest FalconCollared Forest FalconLaughing FalconBat FalconOrange-breasted FalconPlain ChachalacaCrested Guan

Tinamus majorCrypturellus souiCrypturellus cinnamomeusCrypturellus boucardiPelecanus occidentalisPhalacrocorax brasilianusArdea herodiasArdea albaEgretta thulaEgretta caerulea Egretta tricolorBubulcus ibisButorides virescensNycticorax nycticoraxCoragyps atratusCathartes auraSarcoramphus papaDendrocygna autumnalisAnas discorsPandion haliaetusIctinia plumbeaLeucopternis albicollisButeogallus anthracinusButeogallus urubitingaHarpyhaliaetus solitariusButeo magnirostrisHarpia harpyjaSpizaetus tyrannusSpizaetus ornatusMicrastur ruficollisMicrastur semitorquatusHerpetotheres cachinnansFalco rufigularisFalco deiroleucusOrtalis vetulaPenelope purpurascens

PRPRPRPROPRWRWRWRWRWRPRPRWRPRPRPRPRWRWRWRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPR

R

R

PE

SPE

RRAAA

SPEPEA

PERR

AA

SPE

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OMOOOOOOOCMO

OMOOO

CURURRUUCURRRRCCRRRURURRUURRUR-UCRCU

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PUEBLA-OLIVARES ET AL.

APPENDIX 1. Continued.

Common names Species SS CS Records AbundanceGreat CurassowSpotted WoodquailGreater YellowlegsSpotted SandpiperLaughing GullShort-billed PigeonRuddy Ground DoveBlue Ground DoveWhite-tipped DoveGray-fronted DoveGray-chested DoveRuddy Quail DoveOlive-throated ParakeetScarlet MacawBrown-hooded ParrotWhite-crowned ParrotWhite-fronted ParrotMealy ParrotYellow-billed CuckooBlack-billed CuckooSquirrel CuckooStriped CuckooGroove-billed AniVermiculated Screech OwlFerruginous Pygmy OwlMottled OwlShort-tailed NighthawkCommon PauraqueWhite-collared SwiftLong-tailed HermitLittle HermitScaly-breasted HummingbirdWhite-necked JacobinGreen-breasted MangoWhite-bellied EmeraldRufous-tailed HummingbirdBuff-bellied HummingbirdStripe-tailed HummingbirdPurple-crowned FairyLong-billed StarthroatBlack-headed TrogonViolaceous TrogonSlaty-tailed TrogonTody MotmotBlue-crowned Motmot

Crax rubraOdontophorus guttatusTringa melanoleucaActitis maculariaLarus atricillaColumba nigrirostrisColumbina talpacotiClaravis pretiosaLeptotila verreauxiLeptotila rufaxillaLeptotila cassiniGeotrygon montanaAratinga nanaAra macaoPionopsitta haematotisPionus senilisAmazona albifronsAmazona farinosaCoccyzus americanusCoccyzus erythrophthalmusPiaya cayanaTapera naeviaCrotophaga sulcirostrisOtus guatemalaeGlaucidium brasilianumCiccaba virgataLurocalis semitorquatus1

Nyctidromus albicollisStreptoprocne zonarisPhaethornis superciliosusPhaethornis longuemareusPhaeochroa cuvieriiFlorisuga mellivoraAnthracothorax prevostiiAmazilia candidaAmazilia tzacatlAmazilia yucatanensisEupherusa eximiaHeliothryx barrotiHeliomaster longirostrisTrogon melanocephalusTrogon violaceusTrogon massenaHylomanes momotulaMomotus momota

PRPRT

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R

R

RR

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RAA

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AVIFAUNA OF THE YAXCHILÁN NATURAL MONUMENT, MÉXICO

APPENDIX 1. Continued.

Common names Species SS CS Records AbundanceRinged KingfisherAmazon KingfisherGreen KingfisherWhite-necked PuffbirdWhite-whiskered PuffbirdRufous-tailed JacamarCollared AracariKeel-billed ToucanBlack-cheeked WoodpeckerGolden-fronted WoodpeckerSmoky-brown WoodpeckerGolden-olive WoodpeckerChestnut-colored WoodpeckerLineated WoodpeckerPale-billed WoodpeckerRufous-breasted SpinetailBuff-throated Foliage-gleanerPlain XenopsScaly-throated LeaftosserTawny-winged WoodcreeperRuddy WoodcreeperOlivaceous WoodcreeperWedge-billed WoodcreeperNorthern Barred WoodcreeperIvory-billed WoodcreeperStreak-headed WoodcreeperGreat AntshrikeBarred AntshrikePlain AntvireoDot-winged AntwrenDusky AntbirdBare-crowned AntbirdBlack-faced AntthrushScaled AntpittaYellow-bellied TyrannuletGreenish ElaeniaOchre-bellied FlycatcherSepia-capped FlycatcherSlate-headed Tody FlycatcherCommon Tody FlycatcherYellow-olive FlycatcherStub-tailed SpadebillRoyal FlycatcherRuddy-tailed FlycatcherSulphur-rumped Flycatcher

Ceryle torquataChloroceryle amazonaChloroceryle americanaNotharchus macrorhynchosMalacoptila panamensisGalbula ruficaudaPteroglossus torquatusRamphastos sulfuratusMelanerpes pucheraniMelanerpes aurifronsVeniliornis fumigatusPiculus rubiginosusCeleus castaneusDryocopus lineatusCampephilus guatemalensisSynallaxis erythrothoraxAutomolus ochrolaemusXenops minutusSclerurus guatemalensisDendrocincla anabatinaDendrocincla homochroaSittasomus griseicapillusGlyphorhynchus spirurusDendrocolaptes sanctithomaeXiphorhynchus flavigasterLepidocolaptes souleyetiiTaraba majorThamnophilus doliatusDysithamnus mentalisMicrorhopias quixensisCercomacra tyranninaGymnocichla nudicepsFormicarius analisGrallaria guatimalensisOrnithion semiflavumMyiopagis viridicataMionectes oleaginousLeptopogon amaurocephalusPoecilotriccus sylviaTodirostrum cinereumTolmomyias sulphurescensPlatyrinchus cancrominusOnychorhynchus coronatusTerenotriccus erythrurusMyiobius sulphureipygius

PRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPRPROPRPRPRPRPRPRPRPRPRPRPRPRPR

RRRAR

R

ARR

RARARRRR

R

RRR

R

R

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RR

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OMOCOOOO

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OCMOC

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OCMC

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393

PUEBLA-OLIVARES ET AL.

APPENDIX 1. Continued.

Common names Species SS CS Records AbundanceEastern Wood PeweeTropical PeweeYellow-bellied FlycatcherAcadian FlycatcherWillow FlycatcherLeast FlycatcherBright-rumped AttilaRufous MournerDusky-capped FlycatcherGreat Crested FlycatcherGreat KiskadeeBoat-billed FlycatcherSocial FlycatcherSulphur-bellied FlycatcherPiratic FlycatcherTropical KingbirdThrush-like SchiffornisCinnamon BecardRose-throated BecardMasked TityraBlack-crowned TityraRufous PihaLovely CotingaWhite-collared ManakinRed-capped ManakinLoggerhead ShrikeWhite-eyed VireoBlue-headed VireoPhiladelphia VireoYellow-green VireoLesser GreenletTawny-crowned GreenletGreen JayBrown JayNorthern Rough-winged SwallowBarn SwallowMangrove SwallowSpot-breasted WrenWhite-browed WrenWhite-breasted Wood WrenLong-billed GnatwrenSwainson's ThrushWood ThrushClay-colored RobinWhite-throated Robin

Contopus virensContopus cinereusEmpidonax flaviventrisEmpidonax virescensEmpidonax trailliiEmpidonax minimusAttila spadiceusRhytipterna holerythraMyiarchus tuberculiferMyiarchus crinitusPitangus sulphuratusMegarynchus pitanguaMyiozetetes similisMyiodynastes luteiventrisLegatus leucophaiusTyrannus melancholicusSchiffornis turdinusPachyramphus cinnamomeusPachyramphus aglaiaeTityra semifasciataTityra inquisitorLipaugus unirifusCotinga amabilisManacus candeiPipra mentalisLanius ludovicianusVireo griseusVireo solitariusVireo philadelphicusVireo flavoviridisHylophilus decurtatusHylophilus ochraceicepsCyanocorax yncasCyanocorax morioStelgidopteryx serripennisHirundo rusticaTachycineta albilineaThryothorus maculipectusThryothorus albinuchaHenicorhina leucostictaRamphocaenus melanurusCatharus ustulatusHylocichla mustelinaTurdus grayiTurdus assimilis

TPRWRTT

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R

RAR

RR

R

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394

AVIFAUNA OF THE YAXCHILÁN NATURAL MONUMENT, MÉXICO

APPENDIX 1. Continued.

Common names Species SS CS Records AbundanceGray CatbirdBlue-winged WarblerTropical ParulaYellow WarblerChestnut-sided WarblerMagnolia WarblerBlack-throated Green WarblerBay-breasted WarblerBlack-and-white WarblerAmerican RedstartWorm-eating WarblerOvenbirdNorthern WaterthrushKentucky WarblerMacGillivray's WarblerCommon YellowthroatHooded WarblerWilson's WarblerGolden-crowned WarblerYellow-breasted ChatGray-throated ChatBananaquitGray-headed TanagerBlack-throated Shrike TanagerRed-crowned Ant TanagerRed-throated Ant TanagerSummer TanagerCrimson-collared TanagerPasserini's TanagerBlue-gray TanagerYellow-winged TanagerScrub EuphoniaYellow-throated EuphoniaOlive-backed EuphoniaGolden-hooded TanagerGreen HoneycreeperRed-legged HoneycreeperVariable SeedeaterWhite-collared Seedeater Blue SeedeaterOrange-billed SparrowGreen-backed SparrowLincoln's SparrowGrayish SaltatorBuff-throated Saltator

Dumetella carolinensisVermivora pinusParula pitiayumiDendroica petechiaDendroica pensylvanicaDendroica magnoliaDendroica virensDendroica castaneaMniotilta variaSetophaga ruticillaHelmitheros vermivorusSeiurus aurocapillusSeiurus noveboracensisOporornis formosusOporornis tolmieiGeothlypis trichasWilsonia citrinaWilsonia pusillaBasileuterus culicivorusIcteria virensGranatellus sallaeiCoereba flaveolaEucometis penicillataLanio aurantiusHabia rubicaHabia fuscicaudaPiranga rubraRamphocelus sanguinolentusRamphocelus passeriniiThraupis episcopusThraupis abbasEuphonia affinisEuphonia hirundinaceaEuphonia gouldiTangara larvataChlorophanes spizaCyanerpes cyaneusSporophila americanaSporophila torqueolaAmaurospiza concolorArremon aurantiirostrisArremonops chloronotusMelospiza lincolniiSaltator coerulescensSaltator maximus

WRWRPR?WRWRWRWRWRWRWRWRWRWRWRWRWRWRWRPRWRPRPRPRPRPRPRWRPRPRPRPRPRPRPRPRPRPRPRPRPR?PRPRWRPRPR

RR

RRR

A

R

RR

RRR

RR

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OCO

OCMMO

OCMOCMOC

OCMOCMOCM

MOM

OCMOCMOCMOCM

OOCMOCMOM

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OOCO

OCOMOM

OCMO

OCMOMOCOC

OCMMO

OC

CRRURU-RCVUCUC-RUCUURUU-RVUCCUURUUURRCCRCC-UU

395

PUEBLA-OLIVARES ET AL.

APPENDIX 1. Continued.

Common names Species SS CS Records AbundanceBlack-headed SaltatorBlack-faced GrosbeakYellow GrosbeakRose-breasted GrosbeakBlue-black GrosbeakBlue BuntingIndigo BuntingPainted BuntingMelodious BlackbirdGreat-tailed GrackleBlack-cowled OrioleOrchard OrioleSpot-breasted OrioleHooded OrioleYellow-tailed OrioleBaltimore OrioleAltamira OrioleYellow-billed CaciqueChestnut-headed OropendolaMontezuma Oropendola

Saltator atricepsCaryothraustes poliogasterPheucticus chrysopeplusPheucticus ludovicianusCyanocompsa cyanoidesCyanocompsa parellinaPasserina cyaneaPasserina cirisDives divesQuiscalus mexicanusIcterus dominicensisIcterus spuriusIcterus pectoralisIcterus cucullatusIcterus mesomelasIcterus galbulaIcterus gularisAmblycercus holosericeusPsarocolius wagleriPsarocolius montezuma

PRPRPRWRPRPRWRWRPRPRPRWRPRPRPRWRPRPRPRPR

R

A

AR

OCOCO

OMOCMOC

OCMOOO

OMOMOO

OMM

OCMO

OM

CURRURURVRRR-RUU-CUC

1Only recorded by Howell (1989).

396