avifauna of the upper essequibo river and acary … (18) 339-368.pdf · mary forest. our itinerary...

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ORNITOLOGIA NEOTROPICAL 18: 339–368, 2007© The Neotropical Ornithological Society

AVIFAUNA OF THE UPPER ESSEQUIBO RIVER AND ACARY MOUNTAINS, SOUTHERN GUYANA

Mark B. Robbins1, Michael J. Braun2, Christopher M. Milensky3, Brian K. Schmidt3, Waldyke Prince4, Nathan H. Rice5, Davis W. Finch6, & Brian J. O’Shea7

1University of Kansas Natural History Museum & Biodiversity Research Center (KUMNH), 1345 Jayhawk Boulevard, Lawrence, Kansas 66045, USA. E-mail: [email protected]

2Department of Vertebrate Zoology, National Museum of Natural History (USNM), Smithsonian Institution, 4210 Silver Hill Road, Suitland, Maryland 20746, USA.

3Smithsonian Institution, Division of Birds, PO Box 37012, Washington DC, 20013-7012, USA.

4Iwokrama International Centre, PO Box 10630, 77 High Street, Kingston, Georgetown, Guyana.

5Academy of Natural Sciences (ANSP), 1900 Benjamin Franklin Parkway, Philadelphia, Pennsylvania 19103, USA.

691 South Road, East Kingston, New Hampshire 03827, USA.7Department of Biological Sciences and Museum of Natural Science, 119 Foster Hall,

Louisiana State University, Baton Rouge, Louisiana 70803, USA.

Resumen. – Avifauna del alto Río Essequibo y la Sierra de Acary, en el sur de Guyana. –Realizamos inventarios intensivos durante dos temporadas y varias visitas de menor duración en el altoRío Essequibo, en el extremo sur de Guyana, en una zona que está entre las menos impactadaspor humanos en el planeta. En total, registramos 441 especies de aves, incluyendo los primerosregistros de 12 especies para el país. Para otras cuatro especies, colectamos los primeros especimenes delpaís. Presentamos información acerca de abundancia relativa, preferencias de hábitat y estatusreproductivo. La lista de especies para esta región es mayor que la de otro sitio intensamente estudiado alnorte de Manaos, en Brazil, pero menor que la de la Selva Iwokrama en el centro de Guyana. Es probableque las diferencias en la composición de especies entre estos sitios se basen en la heterogeneidad delhábitat.

Abstract. – We conducted two extensive and several shorter-term avifaunal surveys during differentseasons in the upper Essequibo River drainage of extreme southern Guyana, one of theleast human-impacted areas on the planet. A total of 441 avian species were recordedincluding the first Guyana records for 12 species. Four additional species were documented with the firstGuyana specimens. Relative abundance, habitat preferences, and breeding status are presented.The species list for the Upper Essequibo region is higher than that for an intensively studied sitenorth of Manaus, Brazil, but lower than that for the Iwokrama Forest in central Guyana. The likely expla-nation for differences in species composition between these sites is habitat heterogeneity. Accepted 3March 2007.

Key words: Avifauna, Essequibo River, Acary Mountains, Guyana, new records.

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INTRODUCTION

The lowland forests of the Guiana Shield rep-resent one of the largest blocks of pristineforest on earth. The majority of the naturalvegetation remains intact, with estimatesof relatively untouched forest of 80% inGuyana, 90% in Suriname and FrenchGuiana, and 42% in adjacent Brazil (Bryant etal. 1997). The area of southern Guyana,southern Suriname and abutting Brazil hasbeen particularly unaffected by humans.Excluding the Rupununi Savanna, Guyanaforests south of the 4th parallel comprisenearly 7,000,000 ha and have only one perma-nent settlement, the Wai-Wai Amerindianvillage near Gunn’s Strip (D. Clarke pers.observ.; see below). At the extreme southernend of this region, the Wai-Wai own about625,000 ha, these lands being known as theKonashen Community Owned ConservationArea.

From an ornithological standpoint, thisarea remains very poorly known. The nearestthorough avian surveys were conducted about80 km north of Manaus, Amazonas, Brazil(Cohn-Haft et al. 1997) and at the IwokramaForest reserve in central Guyana (Ridgely et al.2005). Oren & Albuquerque’s (1991) updateof Haffer’s (1974) catalog of collecting locali-ties for the Brazilian component of thisregion underscored how few areas have beeneven minimally surveyed, with large sectionsstill totally unknown. The only survey withinthe Guyana portion of this region was madeby Blake (1950) in 1938, without such invalu-able survey tools as mist-nets, tape recorders,and field identification guides.

STUDY AREA AND METHODS

As part of a long-term assessment of thebiodiversity of Guyana, we conducted twoextensive and several shorter surveys duringdifferent seasons in the upper Essequibo

River drainage of extreme southern Guyana.All of our field sites were within theKonashen Community Owned ConservationArea. In August–September 1998 and in Feb-ruary–March 1999, we mounted expeditionsto the headwaters of the Essequibo River.Finch, O’Shea, and others briefly visited thesesame areas in subsequent years. Access to theregion was via small aircraft from George-town to a laterite runway, known as Gunn’sStrip, in an isolated, wet savanna surroundedby thousands of sq. km of contiguous, pri-mary forest. Our itinerary and description ofeach surveyed site follow.

Gunn’s Strip. 01º39’N, 58º37’W, 225 m, Fig. 1.At the time of our work about 200 Wai-Waioccupied a village near the airstrip. Followinghigh water in 2000, the community moved toa point < 1.5 straight-line km upriver. TheGunn’s site is seasonally flooded with muchof the savanna and forest floor inundated formonths, typically from late April through atleast July. We surveyed about 12 km2 of thenatural savanna (Robbins et al. 2005). Inter-spersed within the savanna, which grows ongray, sandy, clay hard pan, are tongues of gal-lery forest and small, isolated patches of low,dense forest known in Guyana as “bushislands”. Moriche or ité palms (Mauritia flexu-osa) were conspicuous at this site. As a resultof periodic burning by the Wai-Wai during thedry season, the savanna-forest interface wasoften abrupt.

This site was surveyed during the follow-ing periods: 16–19 August 1998 & 17–20 Sep-tember 1998 by Robbins, Braun, Milensky,Schmidt, Prince, and Rice; 27 February–8March 1999 & 23–26 March 1999 by Braun,Milensky, Schmidt, and Prince; 18–19November 1999, 28–29 April 2001, 4–6 and22–23 November 2002 by Finch et al, and 4–5and 27–28 October 2006 by O’Shea. No mist-nets were used during the 1998 surveys, Finchet al. and O’Shea visits, but about 10 mist-nets

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were used during the February–March 1999survey.

Sipu River. 01º25’N, 58º57’W, 225 m, Fig. 1.Many downed trees made this narrow water-course difficult to navigate, and where thecamp was established on its south bank, theriver was < 15 m wide. While botanist DavidClarke and our logistical crew cut a trailsouthward toward the Acary (ACK-a-rye)Mountains, we surveyed (15 mist-nets) birdsin the seasonally flooded, swampy forest from21–27 August 1998. Finch et al. worked alongthe Sipu River up to this site from 12–15 &20–21 November 2002. The forest canopywas relatively low and uneven as a result ofpoor soil and the area being flooded for atleast 3 months of the year. All the forest stratawere laden with vines, and the terrain wasvery uneven with hummocks of drier groundintermixed with canal-shaped, low-lying areas.Palms such as Astrocaryum jauari and Euterpeoleracea were a dominant component of theunderstory. During this short period, the riverdropped about 13 cm at our campsite, and bythe time we left the area on 16 September, ithad dropped a total of about 30 cm. Exceptduring infrequent periods of rain, cicadas pro-

duced considerable ambient noise from about07:00 until sunset.

Acary Mountains. 01º23’N, 58º56’W, 225 m, Fig1. This camp was near a stream in tall primaryterra firme forest at the northern base of themountains, about 4 straight-line km from theSipu camp. The area was worked from 28August to 15 September 1998. Finch et al. vis-ited the site from 15–19 November 2002, fol-lowed by O’Shea from 7–18 October 2006. In1998, in order to survey the highest elevations(about 1100 m), we rotated personnel to amini-camp established at 575 m where we setup 3 mist-nets. While two people worked atthe mini-camp, the remaining team surveyedbirds at the base camp, cutting new trails andworking the trail back to the Sipu camp.About 20 mist-nets were used at the basecamp. At about 800 m, the forest appearedwetter with the first sign of large terrestrialferns, moss and bromeliads on the trees. Fre-quent cloud contact along the ridges sup-ported abundant epiphytes on trees andsphagnum moss on the ground, but very fewplants were restricted to the high elevations(D. Clarke pers. observ.). Bamboo (Guadualatifolia) was prevalent along the ridge crests as

FIG. 1. Principal localities mentioned in text.

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a result of tree falls and in the peneplain ofcreeks at the mountain base. A number oftrees were flowering and fruiting from thebase to the crest of the mountains. The high-est summit consisted of granitic bedrock witha thin cover of organic soil and scrub vegeta-tion [Myrtaceae, Mimosa (Fabaceae), andErythroxylon (Erythroxylaceae) (D. Clarke pers.observ.)].

Essequibo River camp. Upriver, about 7 straight-line km from Gunn’s Strip. 01º35 N, 58º38W,250 m, Fig. 1. Surveyed by Braun, Milensky,Schmidt, and Prince from 9 to 22 March1999. Finch et al. worked along the river nearthis site from 6–8 & 21–22 November 2002.A camp was established adjacent to the river.Seasonally flooded, terra firme, and riverinehabitats were surveyed, using 15+ mist-netsnear the base camp. A temporary camp(01º35’N, 58º35’W) was established at thebase of a range of low hills to the east, per-mitting surveys of upper elevations (about700 m) on five mornings. Guyana Lands andSurveys topographical maps refer to thesehills as “Zibingatzako Mountain”. The areabetween this massif and the Essequibo Riveris a mosaic of terra firme and seasonallyflooded forest. The canopy of the terra firmeforest was as high as 40 m, whereas the sea-sonally flooded forest canopy was 20–25 m inheight.

Kamoa River. 01°32’N, 58°50’W, 240 m, Fig. 1.Surveyed by Finch et al. 8–11 November 2002and O’Shea 19–28 October 2006. Both Finchand O’Shea worked from a camp at the abovecoordinates and Finch also worked from asecond camp at 01°31.5’N, 58°46.3’W. DuringO’Shea’s visit, birds were surveyed along trailscut in both directions along the river. Anothertrail was cut perpendicular (north) to theriver, and led to a small rocky hill (about 560m) in terra firme forest. This trail passedthrough an extensive area of low-stature for-

est with many palms, resembling forestaround Gunn’s Strip. Birds were also surveyedby boat on several mornings. Habitat immedi-ately adjacent to the river was seasonallyflooded forest of shorter stature than thatfound along the Sipu or Essequibo rivers; theforest became taller farther away from theriver. The understory of this flooded forestwas very sparse. There appeared to be a highdegree of habitat heterogeneity at this site.

Specimens are deposited at the USNM,KUMNH, and the University of Guyana,Georgetown. Tissue samples are at USNM.Robbins’s and Finch’s recordings have beendeposited and O’Shea’s and Braun’s will bedeposited at the Macaulay Library of NaturalSounds, Cornell University. Taxonomy andnomenclature follow Remsen et al. (2007),except for the Icterus cayanensis complex, wherewe follow Jaramillo & Burke (1999) in orderto make equivalent taxonomic comparisonsamong three sites.

RESULTS AND DISCUSSION

We recorded a total of 441 species in theregion. To our knowledge, the only otherpublished avifaunal information for theUpper Essequibo region is that of Blake(1950) who, in late 1938, amassed nearly 1000skins. Unfortunately, half of this invaluablecollection was lost when his boat capsized inrapids on the return from the Acary Moun-tains and adjacent lowlands. In his assessmentof the region’s affinities, Blake reported thatonly 5 of 125 species that he recorded aboveabout 450 m were subtropical-inhabiting spe-cies: Chestnut-tipped Toucanet (Aulacorhyn-chus derbianus), Golden-olive Woodpecker(Piculus rubiginosus), Strong-billed Wood-creeper (Xiphocolaptes promeropirhynchus),Rufous-rumped Antwren (Terenura callinota),and Sharpbill (Oxyruncus cristatus). We do notconsider either the Piculus or Xiphocolaptes tobe indicative of the subtropical zone, as both

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occur in lowland forests across the GuianaShield, as elsewhere in South America. How-ever, Blake recorded three species whichshould be included in that list as they arefound primarily in montane regions in theGuiana Shield: Collared Trogon (Trogon col-laris), Brown Violetear (Colibri delphinae), andBlackish Pewee (Contopus nigrescens). We docu-mented an additional seven montane speciesthat Blake did not record: VermiculatedScreech-Owl [Megascops (guatemalae) roraimae],Green-bellied Hummingbird [Amazilia (viridi-gaster) cupreicauda], Plain Antvireo (Dysithamnusmentalis), Rufous-tailed Tyrant (Knipolegus poecil-urus), Tepui Greenlet (Hylophilus sclateri),Hepatic Tanager [Piranga (flava) haemalea], andBlue-naped Chlorophonia (Chlorophonia cya-nea). We exclude Cliff Flycatcher (Hirundineaferruginea), which was recorded only alongridge crests, because its preferred habitat ofbare cliff faces is not restricted to mountains.Several species were present in open, scrubbyvegetation along the Acary crests that nor-mally are found in similar habitat in thelowlands, e.g., Red-shouldered Tanager (Tac-hyphonus phoeniceus) and Dusky Antbird (Cerco-macra tyrannina). Two additional species,Tropical Parula (Parula pitiayumi) and Rufous-winged Antwren (Herpsilochmus rufimarginatus),were restricted to the Acary Mountains; bothare found in montane areas as well as locallyin the lowlands in western Guyana, easternVenezuela, and Roraima, Brazil (Hilty 2003,Naka et al. 2006). In Suriname the Herpsiloch-mus is known only from the montane Tafel-berg (O’Shea pers. observ.). Blake recordedtwo species that we failed to detect: GreaterAni (Crotophaga major) and Guira Tanager(Hemithraupis guira). Thus, the species total forthe Upper Essequibo region is at least 443.Additional surveys will probably bring thetotal close to that of the Iwokrama Forest(about 475 species; see below). Most of oursurveys were conducted during the non-breeding season (see below), when relatively

few species were vocalizing, and we undoubt-edly missed several boreal and australmigrants.

During February–March 1999, we conser-vatively documented breeding in 22% (n = 93species) of the avifauna, whereas 20% (n =85) of the avifauna was documented breedingduring August–September 1998 (Appendix 1).Breeding criteria were based on specimengonad data, behavior (displays, carrying nest-ing material), and active nests. In addition, 17species were vocalizing considerably moreduring February–March than during August–September and we considered all of these tobe breeding (Appendix 1). These figuresshould be considered very conservative, asour specimen sample sizes even for commonspecies were low, and many species were notcollected. Because of manpower, our effortwas biased in favor of the 1998 survey (763specimens collected in 1998, 522 specimens in1999). Based on our work elsewhere in Guy-ana (Robbins et al. 2004, unpubl.), the majorityof the Guyana avifauna initiates breeding atthe onset of the main rainy season, typicallyfrom late April into August; exceptionsinclude columbids and other frugivores. Thus,if both our intensive surveys had been con-ducted a month later, i.e., in April and Octo-ber, we predict that we would have found asignificantly higher percentage of the avifaunabreeding, especially in April. Finally, moreNorth American passerine migrants wererecorded during the February to late Marchperiod than in the August to mid-Septemberperiod, when most such migrants have not yetreached Guyana (Appendix 1). An example isthe American Redstart (Setophaga rutilla),which O’Shea found to be fairly common inthe Acary Mountains in mid-October,whereas it was unrecorded there during ourlate August to mid-September survey.

Gunn’s Strip had a fair amount of huntingand burning pressure in the vicinity of the vil-lage, but the other sites were pristine. The

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human population today is a fraction of whatit was when Schomburgk and party visited theregion in 1837. Following the influenza epi-demic of 1918, only a few, scattered villagesremained (Guppy 1958). Indicative of thevery low human presence at the Sipu andAcary sites were the relatively large numbersof Crax and Penelope, seven monkey species,and multiple observations and signs of jaguarand tapir.

Comparison with avifaunas at other Guiana Shieldsites. It is of interest to compare the UpperEssequibo River drainage avifauna with otherextensively inventoried sites to reveal patternsof diversity across the Guiana Shield region.The avian biogeographical affinities of theterra firme forests east of the Rio Branco, Bra-zil are with the Guiana Shield, whereas theavifauna west of this drainage is more closelyrelated to western Amazonia (Naka et al.2006). Thus, we compare the Upper Esse-quibo inventory with the two most completesurveys in this region: 1) The BiologicalDynamics of Forest Fragments Project siteabout 80 km north of Manaus (Cohn-Haft etal. 1997); 2) the Iwokrama Forest reserve incentral Guyana (Ridgely et al. 2005). TheManaus inventory covered a general area ofabout 500 km2 (Fig. 1) and was based on 15years of fieldwork by many experiencedobservers who recorded a total of 394 species.An even higher species list, 476 species, wasrecorded at the Iwokrama Forest (surveyed 8months over 2 years; Ridgely et al. 2005; Fig.1). What explains the diversity differencesamong these three sites? We concur withCohn-Haft et al. (1997) and Naka et al. (2006)that the primary cause of differences in spe-cies richness among sites is habitat heteroge-neity. We believe that the 113 species (112that we found, plus the Hemithraupis thatBlake recorded) recorded in the Upper Esse-quibo region that were not recorded north ofManaus (Appendix 1), are also best explained

by differences in habitat heterogeneitybetween the sites. The Upper Essequibo areaincluded savanna, extensive seasonallyflooded forest, and montane areas in contrastto the largely terra firme Manaus site. In addi-tion to the unique montane element outlinedabove and the obvious contribution ofsavanna to diversity of the Upper Essequiboarea (category 4 under habitat preferences inAppendix 1), a comparison of the species-richThamnophilidae between these two areasunderscores some microhabitat differences.Of the 13 antbird species recorded in theUpper Essequibo area that were absent fromnorth of Manaus, several were restricted tovine tangles in seasonally flooded forest andtwo were highland species (Appendix 1). TheSpot-backed Antwren (Herpsilochmus dorsimacu-latus) was the only thamnophilid found at theManaus area not present at the Upper Esse-quibo region.

Of these three areas, the Iwokrama Foresthas the greatest diversity of habitats. Sixty-seven species were recorded at the UpperEssequibo that were unrecorded fromIwokrama (Appendix 1). Montane andsavanna-inhabiting species, vine tangle spe-cialists, and migrants constitute the bulk ofthe 67 species found in the Upper Essequiboand not Iwokrama. Iwokrama had the uniqueHumiria-dominated low woodland (“MuriScrub”, known as a type of campina in Amazo-nian Brazil) that added six species notrecorded in the Upper Essequibo, and themontane avifauna at Iwokrama included fourtepui species not found in the Acary Moun-tains (Ridgely et al. 2005). Finally, terra firmeforest was more prevalent at Iwokrama thanthe Upper Essequibo.

Cohn-Haft et al. (1997) proposed thatalthough species diversity differed consider-ably among their Manaus locality and twosites in southwestern Amazonia sites, the coreterra firme fauna was surprisingly similarbetween Manaus (n = 264 species) and Manu

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(n = 271 species). Using their same criteria(i.e., treating most of our aerial species as terrafirme-inhabiting) in defining the Upper Esse-quibo terra firme avifauna, we tallied about 248species. Unfortunately, the unpublishedIwokrama Forest species list does not includehabitat preference designations, so terra firmefigures for that locality are not available forcomparison. It should be recognized thatthese numbers should not be considered pre-cise, as habitat requirements of some speciesvary among sites, and not all researchers agreewhen allocating species to particular habitats.Nevertheless, these differences would notdramatically change the overall pattern.

Species accountsWe obtained the first documentation for 12species in Guyana, and the first Guyana speci-mens for four additional species. Detailsand documentation are provided below forall these except Vermiculated Screech-Owl[Megascops (guatemalae) roraimae], VersicoloredEmerald (Amazilia versicolor), Rufous- wingedAntwren (Herpsilochmus rufimarginatus),Rufous-tailed Tyrant (Knipolegus poecilurus) andWhite-naped Seedeater (Dolospingus fringil-loides), details of which we have published ear-lier (Braun et al. 2003; Robbins et al. 2004,2005).

All references to French Guiana, Suri-name, the region north of Manaus, Brazil, andthe Guyana Rupununi are based on informa-tion in Tostain et al. (1992), Haverschmidt &Mees (1994), Cohn-Haft et al. (1997), andRobbins et al. (2004), respectively.

Rusty Tinamou (Crypturellus brevirostris). Thispoorly known tinamou was recorded on onlya few occasions in terra firme forest up to 700m, only at our Acary site. No definite vocal-izations were noted during the 1998 trip, butFinch heard the species in November 2002,and O’Shea obtained a tape recording inOctober 2006. Although these represent the

first records for Guyana, the species is knownfrom French Guiana and northeastern Ama-zonian Brazil, and almost certainly occurs inadjacent southern Suriname. Specimens: 2males, 209 & 233 g (USNM 625102;KUMNH 89676, respectively); female, 295 g(USNM 621695).

Stygian Owl (Asio stygius). An adult male wasflushed during the day on 18 September froma bush island in savanna at Gunn’s Strip byRobbins and Rice (KUMNH 89697) andFinch heard one there on 19 November 1999.Finch et al. heard and recorded two birdsalong the Kuyuwini River (02º 02.85’N, 58º50.40’W; MLNS 106352). These are the firstGuyana records, and we subsequently heardthe species calling from the forest/savannaedge at Kusad Mountain in the Rupununi.Apparently it is still unknown from Suriname,although we predict that ultimately it will befound there, perhaps in the Sipaliwini savannaregion. This species is also known from Ama-zonian savannas (Borges et al. 2001).

Green-bellied Hummingbird (Amazilia viridigastercupreicauda). Robbins and Milensky observedan adult at eye level for about 5 min inexcellent light along an Acary ridge crest at850 m and O’Shea also found this speciesalong a ridge crest in October 2006. Thistaxon, which may deserve species status(Hilty 2003), is more commonly encounteredin montane forests (750–2000 m) of thePantepui in southern Venezuela (Hilty2003), northern Brazil (Naka et al. 2006), andwestern Guyana (Merume Mountains,Quonga, Snyder 1966; Kopinang Mountain,O’Shea pers. observ.). However, there arerecords from as low as 60 m in Venezuela(Hilty 2003), and specimens from lowlandsavanna edge in the southern Rupununi, Guy-ana and Sipaliwini, Suriname. Temporal varia-tion in abundance on Sierra de Lema,Venezuela (Hilty 2003) suggests the possibility

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of seasonal or elevational movement in thisspecies. Rusty-breasted Nunlet (Nonnula rubecula). Anon-breeding female collected in seasonallyflooded forest at our Sipu camp by Schmidtand Braun on 24 August represents the firstGuyana specimen (USNM 625135). It hadbeen observed by Finch et al. on two prioroccasions: 17 April 1996 at Kuyuwini Land-ing (02º05.92’N, 59º14.98’W) and 16 Novem-ber 1997, along the Rewa River (03º44.02’N,58º43.32’W; MLNS 89872). This species hasbeen recorded in Suriname, French Guiana,and across much of Amazonia, includingnorth of Manaus.

Chestnut Woodpecker (Celeus elegans). Blake(1950: 442) remarked that a bird that he col-lected was either an atypical example of thesubspecies hellmayri or was intermediatebetween hellmayri and approximans. The threefemales (USNM 625153; KUMNH 89713-4)that we collected from the Acary Mountainshave pale, creamy yellow crowns with a darkercinnamon color on the elongated crest, brightyellow flanks, and yellow upper tail covertswith varying amounts of cinnamon. Thecrowns of the Acary birds, which closelyresemble nominate elegans, are noticeably palerthan those of birds from northeastern Guy-ana (Berbice River, 05º40’N, 57º53’W,KUMNH 88156; Rockstone, east bank of thelower Essequibo, ANSP 186590), that havebeen assigned to hellmayri.

Rufous-tailed Xenops (Xenops milleri). A breed-ing male (USNM 625165) taken by Braunfrom a mixed-species flock on 27 August atour Sipu camp represents the first Guyanarecord. A few other individuals were seen inmixed-species flocks up to 600 m in theAcary Mountains. It has been recorded at sev-eral localities in Suriname, French Guiana,and north of Manaus.

Spot-throated Woodcreeper (Deconychura sticto-laema). Three non-breeding individuals(USNM 625433-5) were mist-netted in Marchat two of our camps (Appendix 1). These rep-resent the second Guyana locality; the onlyother site is based on an unpublished speci-men of a female collected by A. Capparella on13 August 1994, 5 km NW Mabura Hill(05º19’N, 58º39’W; ANSP 186698). Thiswoodcreeper has recently been observed inSuriname (O’Shea 2005) and it was noted ascommon north of Manaus, as well as presentthroughout Brazilian Amazonia in mixed-spe-cies canopy flocks (M. Cohn-Haft, pers.com.).

Band-tailed Antshrike (Sakesphorus melanotho-rax). O’Shea observed a female in an over-grown clearing in bamboo-dominatedunderstory along a creek near the Acary campon 17 October 2006. The first and only otherGuyana records for this low-density, poorly-known species are those of Finch et al.: pair, 5November 1999 and again 15 April 2001, leftbank of Kuyuwini River (02º05.72’N,59º14.19’W; MLNS 106337, 115727); pair, 21April 2001, left bank of Kassikaityu River(01º50.76’N, 59º04.95’W; MLNS 115771);pair, 22 April 2001, left bank of West Kassika-ityu River (01º50.20’N, 59º12.66’W; MLNS115786); male, 12 November 2001, left bankof Rewa River (03º 06.06’N, 58º38.49’W). Itappears to be rare in Guyana, although it issomewhat more common locally in Suriname(O’Shea & Finch pers. observ.). Spot-winged Antshrike (Pygiptila stellaris). Thisantshrike was uncommon to fairly commonin subcanopy vine tangles of seasonallyflooded forest at our Sipu camp (MLNS131061; 131124). A non-breeding, apparentpair was collected on 7 September (USNM625180, KUMNH 89752) by Schmidt andRobbins. These represent the first Guyanaspecimens. The first Guyana records were

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obtained on 17 April 1997, at Kuyuwini Land-ing (02º05.92’N, 59º14.98’W) and on 14November 1997 along the Rewa River(03º52.15’N, 58º46.41’W; MLNS 89848) byFinch et al. It is known from Suriname andFrench Guiana, but it was not reported fromnorth of Manaus.

Plain Antvireo (Dysithamnus mentalis). Thisspecies was occasionally encountered above850 m at the Acary site. Birds vocalized infre-quently, but a female collected by Schmidt on11 September had recently laid an egg(USNM 625181). These represent the firstdefinite records for Guyana, although it isknown from the Venezuela portion of Mt.Roraima (Phelps 1938, Snyder 1966).Although it has recently been found in Suri-name (O’Shea pers. observ.), it has not beenreported from French Guiana or the Manausregion. Willis’s Antbird (Cercomacra laeta). Thisrange-restricted species was fairly commonalong the forest-savanna interface at Gunn’sStrip (MLNS, n = 5 cuts) and a single bird washeard on 6–8 November 2002 near the Esse-quibo River camp by Finch et al. A series ofspecimens (USNM, KUMNH) indicated nobreeding during August/September or March.These represent the first Guyana records, andwe subsequently recorded it in the extremesouthern portion of the Rupununi savanna.

Olive-green Tyrannulet (Phylloscartes virescens).This inconspicuous, canopy and subcan-opy-dwelling species is poorly known. It wasuncommon in terra firme forest at our Acaryand Essequibo River camps. Presumed pairswere associated with mixed-species flocks. Afemale taken on 9 September had anunshelled egg, 13 x 10 mm, in her ovi-duct (KUMNH 89807). The species is knownfrom scattered localities across the GuianaShield.

Large-headed Flatbill (Ramphotrigon mega-cephalum). O’Shea tape recorded at least twoindividuals in a patch of dense Guadua bam-boo along a creek downstream from theAcary camp on 8 October 2006. This speciesis a bamboo specialist and hence has a patchydistribution across the Amazon basin. Thisrecord extends the species’ known range east-ward ca. 800 km and represents the first forthe Guianas; the nearest published locality isin Amazonas, Venezuela (Hilty 2003).

Pale-bellied Mourner (Rhytipterna immunda).This species was encountered in forest/savanna edge at Gunn’s Strip. Two males fromSeptember and March had small testes(USMN 625235, 622069). Although knownfrom Suriname and French Guiana, these arethe first Guyana records, and we subsequentlyfound it in forest/savanna edge at the extremesouthern edge of the Rupununi savanna.

Boat-billed Tody-Tyrant (Hemitriccus josephi-nae). Species in this genus are inconspicuousand poorly known (Cohn-Haft 2000). Thistody-tyrant is represented by only a handful ofspecimens in museums and very little isknown about it in life. Prior to our work, itwas known in Guyana only from the typespecimen taken in 1910 along the SupenaamRiver near the mouth of the Essequibo River(Chubb 1914). We observed calling males ca.4–9 m above the ground in dense vine tanglesof seasonally flooded forest at our Sipu andEssequibo River camps, and in the southernRupununi. All individuals were in non-breed-ing condition. As described by Donahue(1985), birds gave a “pic-pic”, or sometimes a3-noted series (MLNS 131064; USMN625252, 616573, 622048). This species is nowknown from Suriname, French Guiana, andAmazonian Brazil.

White-eyed Tody-Tyrant (Hemitriccus zoste-rops). At our Acary and Essequibo camps, as

348

ROBBINS ET AL.

many as four individuals/day were encoun-tered 6–10 m above the ground in terra firmeforest. Birds called infrequently (MLNS131059,131105, 131172), although two malescollected in August had enlarged testes. Theserepresent the first Guyana records; the spe-cies is known from Suriname and FrenchGuiana, and was considered common northof Manaus.

Glossy-backed Becard (Pachyramphus surina-mus). An adult male, with a presumed female,was taken out of a canopy mixed-speciesflock at our Sipu camp on 23 August by Rob-bins (KUMNH 89835). It had enlarged, 9 x 5mm, testes. This is the first Guyana record;the species is known from Suriname, FrenchGuiana, and north of Manaus.

White-thighed Swallow (Neochelidon tibialis).Groups of up to 6 individuals were encoun-tered foraging just above the canopy alongridges in the Acary Mountains and at Zibin-gatzako Mountain near our Essequibo Rivercamp. These are the first Guyana records(MLNS 131092). There are only a few recordsfor Suriname, but it is known from FrenchGuiana, and it was considered uncommonnorth of Manaus.

Collared Gnatwren (Microbates collaris). Thiswas a low-density species encountered withina few meters of the ground in terra firme forestfrom 300–750 m. Birds were not singing dur-ing either period and all specimens were innon-breeding condition (USNM, KUNHM).Our specimens are the first for Guyana; thespecies was first recorded in the country inFebruary 1993 in the Kanuku Mountains(Parker et al. 1993). It is also known from sev-eral Suriname and French Guiana localities,and was considered common north ofManaus.

Dotted Tanager (Tangara varia). A non-breed-

ing adult male feeding on melostome berriesat 825 m in the Acary Mountains was col-lected by Robbins on 29 August. This repre-sents the first Guyana specimen (KUMNH89866). The species was encountered on onlyone other occasion at this site. This tanagerhad previously been seen on at least two occa-sions, 15 June 1997, at Turtle Mountain in theIwokrama Reserve (Ridgely et al. 2005) and 19April 2001, along the Kassikaityu River(01º50.51’N, 59º05.16’W) by Finch et al. It isalso known from a few records in Suriname,French Guiana, and north of Manaus.

Epaulet (Icterus cayanensis) & Moriche (Icteruschrysocephalus) Orioles. Although oftentreated as separate species (Meyer de Schau-ensee 1970, Ridgely & Tudor 1989, Jaramillo& Burke 1999), Remsen et al. (2007) treatchrysocephalus as a subspecies of cayanensis.Nonetheless, for consistency of among-sitecomparisons, we follow traditional taxonomyand treat these as separate species. Regardlessof taxonomic treatment, the distribution ofthe two taxa has not been delineated in Guy-ana, as Snyder (1966) made no mention ofcayanensis, stating that chrysocephalus was wide-spread and common. These taxa have beenfound syntopically at our Sipu and Gunn’sStrip sites, Iwokrama Forest (M. Robbins, R.Ridgely pers. observ.), and at Kusad in theRupununi. Although hybridization has beendocumented in Suriname (Haverschmidt &Mees 1994), this has not yet been demon-strated in Guyana.

ACKNOWLEDGMENTS

We especially thank David Clarke for leadingthe logistically difficult 1998 expedition, forsharing his botanical expertise, and for com-menting on the manuscript. Romeo Williams,Claudius Perry, and Wai-Wai Amerindiancommunity were indispensable to the expedi-tions. Vicki Funk and Carol Kelloff of Smith-

349


sonian’s Biological Diversity of the GuianasProgram provided assistance and support.The National Biodiversity Committee of theEnvironmental Protection Agency, the Minis-try of Amerindian Affairs, and the Universityof Guyana provided permission and help forour work. Greg Budney and Mike Andersonprovided catalog data for Robbins’s record-ings at the Macaulay Library of NaturalSounds, Cornell University. Mario Cohn-Haftand Kristof Zyskowski provided valuablecomments on the manuscript. Finch thanksDuane de Freitas for expertly conductingmany trips to remote parts of Guyana. O’Sheathanks the staff of the Rapid Assessment Pro-gram, Center for Applied Biodiversity Sci-ence, Conservation International, as well asCI-Guyana for support during his survey.This is number 119 in the Smithsonian’s Bio-logical Diversity of the Guiana Shield Pro-gram publication series.

REFERENCES

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Braun, M. J., M. B. Robbins, C. M. Milensky, B. J.O’Shea, B. R. Barber, W. Hinds, & W. S. Prince.2003. New birds for Guyana from Mts.Roraima and Ayanganna. Bull. Br. Ornithol.Club 123: 24–33.

Blake, E. R. 1950. Birds of the Acary Mountainssouthern British Guiana. Fieldiana Zool. 32:419–474.

Bryant, D., D. Nielsen, & L. Tangley. 1997. The lastfrontier forests: ecosystems and economies onthe edge. World Resources Institute, Washing-ton, DC.

Chubb, C. 1914. Euscarthmus josephinae. Bull. Br.Ornithol. Club 33:132.

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Haffer, J. 1974. Avian speciation in tropical SouthAmerica. Publ. Nuttall Ornithol. Club 14: 1–390.

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Meyer de Schauensee, R. 1970. A guide to the birdsof South America. Livingston Publ. Co., Wyn-newood, Pennsylvania.

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O’Shea, B. J. 2005. Notes on birds of the Sipaliwinisavanna and other localities in southern Suri-name, with six new species for the country.Ornitol. Neotrop. 16: 361–370.

Parker, T. A., III, R. B. Foster, L. H. Emmons, P.Freed, A. B. Forsyth, & B. D. Gill. 1993. A bio-logical assessment of the Kanuku MountainRegion of southwestern Guyana. Rapid Assess-ment Program Paper 5, Conservation Interna-tional, Washington, DC.

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Remsen, J. V., A. Jaramillo, M. Nores, J. F. Pacheco,M. B. Robbins, T. S. Schulenberg, F. G. Stiles, J.M. C. da Silva, D. F. Stotz, & K. J. Zimmer.

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2007. A classification of the bird species ofSouth America. American Ornithologists’Union. http//www.museum.lsu.edu/~Rem-sen/SACCBaseline.html

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Robbins, M.B., M. J. Braun, C. J. Huddleston, D. W.Finch, & C. M. Milensky. 2005. Systematics,natural history, and first Guyana records of theWhite-naped Seedeater (Dolospingus fringilloides).Ibis 147: 334–341.

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351

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APPENDIX 1. Relative abundance, habitat preferences, and breeding status for the Upper Essequibo River and Acary Mountain avifauna.

English names Scientific names Sipu/Acary1 Gunn’s Strip1

Essequibo River1

Kamoa1 Unique to Upper Essequibo Region2

Habitat3 Breeding status4

TinamousGreat TinamouCinereous TinamouLittle TinamouUndulated TinamouRed-legged TinamouVariegated TinamouRusty Tinamou

Ducks, GeeseMuscovy Duck

Curassows, GuansVariable ChachalacaMarail GuanBlue-throated Piping-GuanBlack Curassow

QuailsMarbled Wood-Quail

CormorantsNeotropic Cormorant

AnhingasAnhinga

HeronsZigzag HeronRufescent Tiger-HeronAgami HeronBoat-billed HeronBlack-crowned Night-HeronStriated HeronCattle Egret

TinamidaeTinamus majorCrypturellus cinereusCrypturellus souiCrypturellus undulatusCrypturellus erythropsCrypturellus variegatusCrypturellus brevirostris

AnatidaeCairina moschata

CracidaeOrtalis motmotPenelope marailPipile cumanensisCrax alector

OdontophoridaeOdontophorus gujanensis

PhalacrocoracidaePhalacrocorax brasilianus

AnhingidaeAnhinga anhinga

ArdeidaeZebrilus undulatusTigrisoma lineatumAgamia agamiCochlearius cochleariusNycticorax nycticoraxButorides striataBubulcus ibis

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L.APPENDIX 1. Continued.


Essequibo River1



Capped HeronCocoi HeronGreat EgretLittle Blue Heron

IbisesGreen Ibis

VulturesLesser Yellow-headed VultureGreater Yellow-headed VultureBlack VultureKing Vulture

OspreysOsprey

Hawks, EaglesHook-billed KiteSwallow-tailed KiteDouble-toothed KitePlumbeous KiteBicolored HawkWhite HawkBlack-faced HawkGreat Black-HawkRoadside HawkShort-tailed HawkWhite-tailed HawkHarpy EagleBlack-and-white Hawk-EagleBlack Hawk-EagleOrnate Hawk-Eagle

Pilherodius pileatusArdea cocoiArdea albaEgretta caerulea

ThreskiornithidaeMesembrinibis cayennensis

CathartidaeCathartes burrovianusCathartes melambrotusCoragyps atratusSarcoramphus papa

PandionidaePandion haliaetus

AccipitridaeChondrohierax uncinatusElanoides forficatusHarpagus bidentatusIctinia plumbeaAccipiter bicolorLeucopternis albicollisLeucopternis melanopsButeogallus urubitingaButeo magnirostrisButeo brachyurusButeo albicaudatusHarpia harpyjaSpizaetus melanoleucusSpizaetus tyrannusSpizaetus ornatus

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APPENDIX 1. Continued.


Essequibo River1



Falcons, CaracarasBlack CaracaraRed-throated CaracaraYellow-headed CaracaraLaughing FalconBarred Forest-FalconLined Forest-FalconSlaty-backed Forest-FalconCollared Forest-FalconBat Falcon

TrumpetersGray-winged Trumpeter

RailsGray-breasted CrakeAsh-throated CrakeGray-necked Wood-RailAzure Gallinule

SungrebesSungrebe

SunbitternsSunbittern

PloversAmerican Golden-Plover

SandpipersLesser YellowlegsSolitary SandpiperSpotted SandpiperUpland SandpiperWhite-rumped Sandpiper

FalconidaeDaptrius aterIbycter americanusMilvago chimachimaHerpetotheres cachinnansMicrastur ruficollisMicrastur gilvicollisMicrastur mirandolleiMicrastur semitorquatusFalco rufigularis

PsophiidaePsophia crepitans

RallidaeLaterallus exilisPorzana albicollisAramides cajaneaPorphyrio flavirostris

HeliornithidaeHeliornis fulica

EurypygidaeEurypyga helias

CharadriidaePluvialis dominica

ScolopacidaeTringa flavipesTringa solitariaActitis maculariusBartramia longicaudaCalidris fuscicollis

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Essequibo River1



Pectoral SandpiperSouth American Snipe

TernsYellow-billed Tern

Pigeons, DovesCommon Ground-DoveRuddy Ground-DoveBlue Ground-DoveScaled PigeonPale-vented PigeonPlumbeous PigeonRuddy PigeonGray-fronted DoveRuddy Quail-Dove

ParrotsBlue-and-yellow MacawScarlet MacawRed-and-green MacawChestnut-fronted MacawRed-bellied MacawWhite-eyed ParakeetPainted ParakeetGolden-winged ParakeetSapphire-rumped ParrotletBlack-headed ParrotCaica ParrotBlue-headed ParrotDusky ParrotBlue-cheeked Parrot

Calidris melanotosGallinago paraguaiae

LaridaeSternula superciliaris

ColumbidaeColumbina passerinaColumbina talpacotiClaravis pretiosaPatagioenas speciosaPatagioenas cayennensisPatagioenas plumbeaPatagioenas subvinaceaLeptotila rufaxillaGeotrygon montana

PsittacidaeAra araraunaAra macaoAra chloropterusAra severusOrthopsittaca manilataAratinga leucopthalmusPyrrhura pictaBrotogeris chrysopterusTouit purpuratusPionites melanocephalusPionopsitta caicaPionus menstruusPionus fuscusAmazona dufresniana

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Orange-winged ParrotRed-fan Parrot

HoatzinsHoatzin

CuckoosLittle CuckooSquirrel CuckooBlack-bellied CuckooPavonine CuckooSmooth-billed Ani

Typical OwlsTawny-bellied Screech-OwlVermiculated Screech-OwlCrested OwlSpectacled OwlAmazonian Pygmy-OwlStygian Owl

PotoosLong-tailed PotooCommon PotooWhite-winged PotooRufous Potoo

Nighthawks, NightjarsSemicollared NighthawkCommon PauraqueBlackish NightjarLadder-tailed Nightjar

SwiftsSwift sp.

Amazona amazonicaDeroptyus accipitrinus

OpisthocomidaeOpisthocomus hoazin

CuculidaeCoccycua minutaPiaya cayanaPiaya melanogasterDromococcyx pavoninusCrotophaga ani

StrigidaeMegascops watsoniiMegascops guatemalaeLophostrix cristataPulsatrix perspicillataGlaucidium hardyiAsio stygius

NyctibiidaeNyctibius aethereusNyctibius griseusNyctibius leucopterusNyctibius bracteatus

CaprimulgidaeLurocalis semitorquatusNyctidromus albicollisCaprimulgus nigrescensHydropsalis climacocerca

ApodidaeCypseloides sp.

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White-collared SwiftBand-rumped SwiftGray-rumped SwiftChapman's SwiftShort-tailed SwiftWhite-tipped SwiftFork-tailed Palm-SwiftLesser Swallow-tailed Swift

HummingbirdsRufous-breasted HermitPale-tailed BarbthroatEastern Long-tailed HermitStraight-billed HermitReddish HermitGray-breasted SabrewingWhite-necked JacobinBrown VioletearCrimson TopazTufted CoquetteRacket-tailed CoquetteFork-tailed WoodnymphWhite-chinned HummingbirdRufous-throated HummingbirdGreen-tailed GoldenthroatVersicolored HummingbirdGreen-bellied HummingbirdBlack-eared FairyAmethyst Woodstar

Trogons

Streptoprocne zonarisChaetura spinicaudusChaetura cinereiventrisChaetura chapmaniChaetura brachyuraAeronautes montivagusTachornis squamataPanyptila cayennensis

TrochilidaeGlaucis hirsutusThrenetes leucurusPhaethornis superciliosusPhaethornis bourcieriPhaethornis ruberCampylopterus largipennisFlorisuga mellivoraColibri delphinaeTopaza pellaLophornis ornatusDiscosura longicaudaThalurania furcataHylocharis cyanusHylocharis sapphirinaPolytmus theresiaeAmazilia versicolorAmazilia viridigasterHeliothryx auritaCalliphlox amethystina

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KingfishersRinged KingfisherAmazon KingfisherGreen KingfisherGreen-and-rufous KingfisherPygmy Kingfisher

MotmotsBlue-crowned Motmot

JacamarsYellow-billed JacamarGreen-tailed JacamarBronzy JacamarParadise JacamarGreat Jacamar

PuffbirdsGuianan PuffbirdPied PuffbirdSpotted PuffbirdCollared PuffbirdWhite-chested PuffbirdRusty-breasted NunletBlack NunbirdSwallow-winged Puffbird

Barbets

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AlcedinidaeMegaceryle torquataChloroceryle amazonaChloroceryle americanaChloroceryle indaChloroceryle aenea

MomotidaeMomotus momota

GalbulidaeGalbula albirostrisGalbula galbulaGalbula leucogastraGalbula deaJacamerops aureus

BucconidaeNotharchus macrorhynchusNotharchus tectusBucco tamatiaBucco capensisMalacoptila fuscaNonnula rubeculaMonasa atraChelidoptera tenebrosa

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WoodpeckersGolden-spangled PiculetYellow-throated WoodpeckerGolden-green WoodpeckerGolden-olive WoodpeckerChestnut WoodpeckerWaved WoodpeckerCream-colored WoodpeckerRinged WoodpeckerLineated WoodpeckerGolden-collared WoodpeckerRed-necked WoodpeckerCrimson-crested Woodpecker

OvenbirdsRuddy SpinetailYellow-chinned SpinetailPoint-tailed PalmcreeperCinnamon-rumped Foliage-gleanerRufous-tailed Foliage-gleanerRufous-rumped Foliage-gleanerOlive-backed Foliage-gleaner

Capito nigerRamphastidae

Aulacorhynchus derbianusSelenidera culikPteroglossus aracariPteroglossus viridisRamphastos vitellinusRamphastos tucanus

PicidaePicumnus exilisPiculus flavigulaPiculus chrysochlorosPiculus rubiginosusCeleus elegansCeleus undatusCeleus flavusCeleus torquatusDryocopus lineatusVeniliornis cassiniCampephilus rubricollisCampephilus melanoleucos

FurnariidaeSynallaxis rutilansCerthiaxis cinnamomeusBerlepschia rikeriPhilydor pyrrhodesPhilydor ruficaudatumPhilydor erythrocercumAutomolus infuscatus

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Striped WoodcreeperChestnut-rumped WoodcreeperBuff-throated WoodcreeperLineated WoodcreeperCurve-billed Scythebill

Typical AntbirdsFasciated AntshrikeBlack-throated AntshrikeGreat AntshrikeBlack-crested Antshrike

Automolus ochrolaemusAutomolus rufipileatusAutomolus rubiginosusXenops minutusXenops milleriSclerurus rufigularisSclerurus caudacutusDendrocincla fuliginosaDendrocincla merulaDeconychura stictolaemaGlyphorynchus spirurusSittasomus griseicapillusDendrexetastes rufigulaHylexetastes perrotiiDendrocolaptes certhiaDendrocolaptes picumnusXiphocolaptes promeropi-rhynchusXiphorhynchus obsoletusXiphorhynchus pardalotusXiphorhynchus guttatusLepidocolaptes albolineatusCampylorhamphus procur-voides

ThamnophilidaeCymbilaimus lineatusFrederickena viridisTaraba majorSakesphorus canadensis

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Band-tailed AntshrikeMouse-colored AntshrikeGuianan Slaty-AntshrikeAmazonian AntshrikeSpot-winged AntshrikePlain AntvireoDusky-throated AntshrikeCinereous AntshrikePygmy AntwrenGuianan Streaked-AntwrenRufous-bellied AntwrenBrown-bellied AntwrenWhite-flanked AntwrenLong-winged AntwrenGray AntwrenSpot-tailed AntwrenTodd's Antwren

Rufous-winged Antwren

Dot-winged AntwrenRufous-rumped AntwrenAsh-winged AntwrenGray AntbirdDusky AntbirdWillis's AntbirdWhite-browed AntbirdWarbling AntbirdBlack-chinned Antbird

Sakesphorus melanothoraxThamnophilus murinusThamnophilus punctatusThamnophilus amazonicusPygiptila stellarisDysithamnus mentalisThamnomanes ardesiacusThamnomanes caesiusMyrmotherula brachyuraMyrmotherula surinamensisMyrmotherula guttataMyrmotherula gutturalisMyrmotherula axillarisMyrmotherula longipennisMyrmotherula menetriesiiHerpsilochmus sticturusHerpsilochmus stictocepha-lusHerpsilochmus rufimargina-tusMicrorhopias quixensisTerenura callinotaTerenura spodioptilaCercomacra cinerascensCercomacra tyranninaCercomacra laetaMyrmoborus leucophrysHypocnemis cantatorHypocnemoides melanopogon

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1,2,31,21,21,21,2

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3,5,13,43

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Silvered AntbirdSpot-backed AntbirdScale-backed AntbirdBlack-headed AntbirdSpot-winged AntbirdFerruginous-backed AntbirdBlack-throated AntbirdWing-banded AntbirdWhite-plumed AntbirdRufous-throated Antbird

Ground AntbirdsRufous-capped AntthrushBlack-faced AntthrushVariegated AntpittaSpotted AntpittaThrush-like Antpitta

GnateatersChestnut-belted Gnateater

Tyrant FlycatchersSooty-headed TyrannuletYellow-crowned TyrannuletGray ElaeniaForest ElaeniaYellow-crowned ElaeniaYellow-bellied ElaeniaLesser ElaeniaPlain-crested ElaeniaRufous-crowned ElaeniaWhite-lored Tyrannulet

Sclateria naeviaHylophylax naeviusHylophylax poecilinotusPercnostola rufifronsPercnostola leucostigmaMyrmeciza ferrugineaMyrmeciza atrothoraxMyrmornis torquataPithys albifronsGymnopithys rufigula

FormicariidaeFormicarius colmaFormicarius analisGrallaria variaHylopezus maculariusMyrmothera campanisona

ConopophagidaeConopophaga aurita

TyrannidaePhyllomyias griseicepsTyrannulus elatusMyiopagis canicepsMyiopagis gaimardiiMyiopagis flavivertexElaenia flavogasterElaenia chiriquensisElaenia cristataElaenia ruficepsOrnithion inerme

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Essequibo River1



Southern Beardless-TyrannuletRinged AntpipitSlender-footed TyrannuletOlive-green TyrannuletOchre-bellied FlycatcherMcConnell's FlycatcherSepia-capped FlycatcherAmazonian Scrub-FlycatcherShort-tailed Pygmy-TyrantDouble-banded Pygmy-TyrantHelmeted Pygmy-TyrantBoat-billed Tody-TyrantWhite-eyed Tody-TyrantCommon Tody-FlycatcherPainted Tody-FlycatcherOlivaceous FlatbillYellow-margined FlatbillGray-crowned FlatbillOchre-lored FlatbillWhite-crested SpadebillCinnamon-crested SpadebillGolden-crowned SpadebillAmazonian Royal-FlycatcherWhiskered FlycatcherRuddy-tailed FlycatcherCinnamon Manakin-TyrantCliff FlycatcherEuler's FlycatcherFuscous Flycatcher

Camptostoma obsoletumCorythopis torquatusZimmerius gracilipesPhylloscartes virescensMionectes oleagineusMionectes macconnelliLeptopogon amaurocephalusSublegatus obscuriorMyiornis ecaudatusLophotriccus vitiosusLophotriccus galeatusHemitriccus josephinaeHemitriccus zosteropsTodirostrum cinereumTodirostrum pictumRhynchocyclus olivaceusTolmomyias assimilisTolmomyias poliocephalusTolmomyias flaviventrisPlatyrinchus platyrhynchosPlatyrinchus saturatusPlatyrinchus coronatusOnychorhynchus coronatusMyiobius barbatusTerenotriccus erythrurusNeopipo cinnamomeaHirundinea ferrugineaLathrotriccus euleriCnemotriccus fuscatus

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Tropical PeweeBlackish PeweeAmazonian Black-TyrantRufous-tailed TyrantDrab Water-TyrantPiratic FlycatcherRusty-margined FlycatcherDusky-chested FlycatcherGreat KiskadeeYellow-throated FlycatcherBoat-billed FlycatcherSulphury FlycatcherWhite-throated KingbirdTropical KingbirdFork-tailed FlycatcherGrayish MournerPale-bellied MournerDusky-capped FlycatcherSwainson's FlycatcherShort-crested FlycatcherLarge-headed Flatbill

Rufous-tailed FlatbillCinnamon AttilaBright-rumped Attila

SharpbillsSharpbill

CotingasSpangled Cotinga

Contopus cinereusContopus nigrescensKnipolegus poecilocercusKnipolegus poecilurusOchthornis littoralisLegatus leucophaiusMyiozetetes cayanensisMyiozetetes luteiventrisPitangus sulphuratusConopias parvusMegarynchus pitanguaTyrannopsis sulphureaTyrannus albogularisTyrannus melancholicusTyrannus savanaRhytipterna simplexRhytipterna immundaMyiarchus tuberculiferMyiarchus swainsoniMyiarchus feroxRamphotrigon megacepha-lumRamphotrigon ruficaudaAttila cinnamomeusAttila spadiceus

OxyruncidaeOxyruncus cristatus

CotingidaeCotinga cayana

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Essequibo River1



Pompadour CotingaWhite BellbirdScreaming PihaPurple-throated FruitcrowCrimson FruitcrowCapuchinbirdGuianan Red-CotingaGuianan Cock-of-the-Rock

ManakinsSaffron-crested ManakinTiny Tyrant-ManakinBlack ManakinWhite-bearded ManakinWhite-throated ManakinGolden-headed ManakinWhite-crowned ManakinWhite-fronted Manakin

Incertae SedisBlack-tailed TityraThrush-like SchiffornisWing-barred PipritesCinereous MournerGlossy-backed BecardWhite-winged BecardBlack-capped BecardCinereous BecardPink-throated Becard

VireosRufous-browed Peppershrike

Xipholena puniceaProcnias albusLipaugus vociferansQuerula purpurataHaematoderus militarisPerissocephalus tricolorPhoenicircus carnifexRupicola rupicola

PipridaeNeopelma chrysocephalusTyranneutes virescensXenopipo atronitensManacus manacusCorapipo gutturalisPipra erythrocephalaPipra pipraLepidothrix serena

Tityra cayanaSchiffornis turdinaPiprites chlorisLaniocera hypopyrraPachyramphus surinamusPachyramphus polychopterusPachyramphus marginatusPachyramphus rufusPachyramphus minor

VireonidaeCyclarhis gujanensis

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Slaty-capped Shrike-VireoRed-eyed VireoLemon-chested GreenletTepui GreenletBuff-cheeked GreenletTawny-crowned Greenlet

JaysCayenne Jay

SwallowsWhite-winged SwallowBrown-chested MartinGray-breasted MartinWhite-banded SwallowBlack-collared SwallowWhite-thighed SwallowBank SwallowBarn Swallow

WrensCoraya WrenSouthern House-WrenWhite-breasted Wood-WrenMusician WrenWing-banded Wren

Gnatwrens, GnatcatchersCollared GnatwrenLong-billed GnatwrenTropical GnatcatcherGuianan Gnatcatcher

Thrushes

Vireolanius leucotisVireo olivaceusHylophilus thoracicusHylophilus sclateriHylophilus muscicapinusHylophilus ochraceiceps

CorvidaeCyanocorax cayanus

HirundinidaeTachycineta albiventerProgne taperaProgne chalybeaAtticora fasciataAtticora melanoleucaNeochelidon tibialisRiparia ripariaHirundo rustica

TroglodytidaeThryothorus corayaTroglodytes musculusHenicorhina leucostictaCyphorhinus aradaMicrocerculus bambla

PolioptilidaeMicrobates collarisRamphocaenus melanurusPolioptila plumbeaPolioptila guianensis

Turdidae

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54,54,5551

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2,3,5,14,32,12,12,1

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VeeryGray-cheeked ThrushBlack-billed ThrushCocoa ThrushWhite-necked Thrush

TanagersBlack-faced TanagerRed-billed Pied TanagerFulvous Shrike-TanagerFlame-crested TanagerFulvous-crested TanagerWhite-shouldered TanagerRed-shouldered TanagerSilver-beaked TanagerBlue-gray TanagerPalm TanagerBlue-backed TanagerTurquoise TanagerOpal-rumped TanagerParadise TanagerSpotted TanagerDotted TanagerBay-headed TanagerBlue DacnisBlack-faced DacnisShort-billed HoneycreeperPurple HoneycreeperRed-legged HoneycreeperGreen Honeycreeper

Catharus fuscescensCatharus minimusTurdus ignobilisTurdus fumigatusTurdus albicollis

ThraupidaeSchistochlamys melanopisLamprospiza melanoleucaLanio fulvusTachyphonus cristatusTachyphonus surinamusTachyphonus luctuosusTachyphonus phoeniciusRamphocelus carboThraupis episcopusThraupis palmarumCyanicterus cyanicterusTangara mexicanaTangara veliaTangara chilensisTangara punctataTangara variaTangara gyrolaDacnis cayanaDacnis lineataCyanerpes nitidusCyanerpes caeruleusCyanerpes cyaneusChlorophanes spiza

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Yellow-backed TanagerSwallow-Tanager

Incertae SedisHepatic TanagerSummer TanagerBananaquit

Emberizine FinchesBlue-black GrassquitYellow-bellied SeedeaterChestnut-bellied Seed-FinchWhite-naped SeedeaterPectoral SparrowRed-capped Cardinal

Grosbeaks, SaltatorsBuff-throated SaltatorSlate-colored GrosbeakBlue-black GrosbeakYellow-green Grosbeak

Wood WarblersTropical ParulaBlackpoll WarblerAmerican RedstartRiver Warbler

Incertae SedisRose-breasted ChatNew World BlackbirdsGreen OropendolaCrested OropendolaYellow-rumped Cacique

Hemithraupis flavicollisTersina viridis

Piranga flavaPiranga rubraCoereba flaveola

EmberizidaeVolatinia jacarinaSporophila nigricollisOryzoborus angolensisDolospingus fringilloidesArremon taciturnusParoaria gularis

CardinalinaeSaltator maximusSaltator grossusCyanocompsa cyanoidesCaryothraustes canadensis

ParulidaeParula pitiayumiDendroica striataSetophaga ruticillaPhaeothlypis rivularis

Granatellus pelzelniIcteridaePsarocolius viridisPsarocolius decumanusCacicus cela

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Red-rumped CaciqueEpaulet OrioleMoriche OrioleGiant CowbirdRed-breasted BlackbirdBobolink

Cardueline FinchesPlumbeous EuphoniaViolaceous EuphoniaGolden-bellied EuphoniaWhite-vented EuphoniaGolden-sided EuphoniaBlue-naped Chlorophonia

Cacicus haemorrhousIcterus cayanensisIcterus chrysocephalusMolothrus oryzivorusSturnella militarisDolichonyx oryzivorus

FringillidaeEuphonia plumbeaEuphonia violaceaEuphonia chrysopastaEuphonia minutaEuphonia cayennensisChlorophonia cyanea

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XS

S

UX

U

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U

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M

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44

1322

2,11

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*

*

1Relative abundance: C = common; more than 20 individuals/day; F = fairly common; 5-20 individuals/day; U = uncommon; present in small numbers (<5 individuals/day); not encountered daily even in prime habitat; S = scarce; only occasionally encountered in small numbers; X= single record; h = onlyabove 300 m.

2Species absent from either Manaus (M) and/or Iwokrama (I).

3Habitat: 1= terra firme forest, 2 = seasonally flooded forest, 3 = forest edge, clearings, 4 = savannah (includes grassland and bush islands), 5 = river, riveredge, 6= aerial.

4Asterisk indicates breeding during February/March; # indicates breeding during Aug/Sept period; V=species vocalizing more in Feb/March vs Aug/Sept.

avifauna of the upper essequibo river and acary … (18) 339-368.pdf · mary forest. our itinerary...

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