avifauna of the guyana southern rupununi...

ORNITOLOGIA NEOTROPICAL 15: 173–200, 2004© The Neotropical Ornithological Society

AVIFAUNA OF THE GUYANA SOUTHERN RUPUNUNI, WITH COMPARISONS TO OTHER SAVANNAS OF NORTHERN SOUTH

AMERICA

Mark B. Robbins1, Michael J. Braun2 & Davis W. Finch3

1Division of Birds, University of Kansas Natural History Museum, Lawrence, KS 66045, USA. E-mail: [email protected]

2Department of Systematic Biology & Laboratory of Analytical Biology, National Museum of Natural History, Smithsonian Institution, 4210 Silver Hill Rd., Suitland, Maryland 20746, USA.

3WINGS, 1643 North Alvernon Way, Suite 105, Tucson, AZ 85712, USA.

Resumen. – Avifauna del Rupununi del Sur, Guyana, con comparaciones con otras sabanas delnorte de Sudamérica. – Inventarios en cinco sitios de la sabana del Rupununi del Sur en Guyanageneraron gran cantidad de información acerca de la composición, abundancia relativa y estatusreproductivo de la avifauna de esta región tan poco conocida. Registros notables de esos inventariosincluyeron el descubrimiento de una población del Cardenalito de Venezuela (Carduelis cucullata), y unaespecie críptica no descrita de atrapamoscas. La comparación de nuestros resultados con listas de especiesde las principales sabanas en el norte de Sudamérica apoyan la idea de que la avifauna de Roraima-Rupununi es más similar a la de la contigua gran sabana de Venezuela, mientras que la avifauna de lasabana Sipaliwini del sur de Surinam es más similar a la del Amapá del noreste de Brasil. Se presentancomentarios detallados para 30 especies, de las cuales 10 son nuevas para Guyana.

Abstract. – Surveys of five sites in the southern Rupununi savanna of Guyana contribute considerably toour knowledge of the species composition, relative abundance, and breeding status of the avifauna of thispoorly known region. Highlights from these surveys include the discovery of a previously unknown popu-lation of the endangered Red Siskin (Carduelis cucullata), and an unrecognized cryptic species of flycatcher.Comparison of our Rupununi results with species lists for several major savannas in northern South Amer-ica further supports the conclusion that the avifauna of the Roraima-Rupununi is most similar to the con-tiguous “Gran Sabana” of Venezuela, whereas the Sipaliwini savanna avifauna of southern Suriname ismost similar to the Amapá of northeastern Brazil. Details are presented for 30 species, including 10 newfor Guyana. Accepted 18 October 2003.

Key words: Avifauna, biogeography, savanna, Guyana, southern Rupununi.

INTRODUCTION

The status and distribution of Neotropicalavifaunas in savanna woodland and savannagrassland are considered to be well under-stood because of the relatively low speciesdiversity coupled with the accessibility ofthese habitats (Silva 1995a, 1995b, Stotz et al.

1996). Nevertheless, very little has been pub-lished on the avifauna of the extensiveRoraima-Rupununi savannas which cover c.54,000 km2 of extreme northern Brazil andsouthwestern Guyana (Fig. 1). Aside fromPinto’s (1966) avifaunal catalog of the Brazil-ian territory of Roraima and the work ofMoskovits et al. (1985) and Silva (1998) on the

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Ilha de Maracá, only general comments onsavanna birds in this region are included inSnyder (1966) and Sick (1993). Oren & Albu-querque (1991) identified much of theRoraima savanna in Brazil as one of the high-est priority areas in need of new ornithologi-cal collections. Indeed, their update ofHaffer’s (1974) catalog of collecting localitiesfor this region of Brazil did not include a sin-gle new site.

The adjoining Rupununi savanna of Guy-ana is divided into northern and southernsubregions by the western extension of theKanuku Mountains (Figs 1 and 2). Virtuallyall ornithological collecting localities are inthe north Rupununi (see p. 97 in Stephens &Traylor 1985), and only a few of these locali-ties are represented by more than a handful of19th century specimens. The only published

avifaunal work on the south Rupununi is thatof Mees & Mees-Balchin (1990) and Mees(2000), who spent a total of about threemonths in the region in 1989 and 1992.

The savanna woodland/grassland com-munity is one of the most impacted biomes inSouth America (Silva 1995c, Stotz et al. 1996).Fortunately, the Roraima-Rupununi regionhas not yet experienced the devastating con-version of grassland to cropland that hasoccurred over such broad areas of Brazil(Cavalcanti 1988, Klink et al. 1993). However,the extensive, annual use of fire during thedry season has undoubtedly had a negativeimpact on a number of grassland and forestedge species in this region.

Here we present results from five inten-sively surveyed sites in the southern Rupu-nuni of Guyana, list noteworthy observations

FIG. 1. Regional map of northern South America with major savanna regions shaded. Inset quadrangleoutlines map of Figure 2. Smaller coastal savannas also exist in northern Guyana, Suriname, French Gui-ana and on Marajó Island at the mouth of the Amazon (not shown). Maps drawn with ArcMap 8.2 (ESRICorp. 2002). Savanna distributions from World Wildlife Fund Ecoregions data, available in ArcMap.

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of several species elsewhere in the GuyanaRupununi, and compare the southern Rupu-nuni with other savannas of northern SouthAmerica. Highlights include the discovery of apreviously unknown population of the endan-gered Red Siskin (Carduelis cucullata; Robbins etal. 2003), and an unrecognized cryptic speciesof flycatcher (in prep.). These discoveriesalong with a number of other surprises, con-tribute considerably to the ornithologicalinterest of this region.

STUDY AREA AND METHODS

The Rupununi savanna of southwestern Guy-ana is a vast low-lying plain, 100–200 m in ele-vation and c. 13,000 km2 in extent (Hills

1976). The savanna extends for many kilome-ters to the west in the Brazilian state ofRoraima, where it is known as the Rio Brancosavanna. Indeed, the total area of savannathere is considerably greater than in Guyana(Eden 1964, Cole 1986). The rolling plains arecovered by grasses, dominated by the peren-nial bunch grass Trachypogon plumosus(Poaceae), interspersed with the dominantsavanna tree, Curatella americana (Dilleniaceae;known in Guyana by the Brazilian name“caimbé” or as “sandpaper tree”). Scatteredthroughout the southern Rupununi, and con-tinuing in adjacent Brazil, are isolated granitemountains that range from small dome-likehills with sparse semi-humid vegetation to rel-atively tall mountains such as Kusad (800 m)

FIG. 2. Map of study area in southwestern Guyana with savannas shaded light gray, mountains dark gray.Triangles indicate main study sites. Circles mark other localities mentioned in text. Savanna distributionsfrom World Wildlife Fund Ecoregions data, available in Huber et al. (1995).

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that cover several square kilometers withmore mesic forest. Within the savanna areisolated patches of forest ranging in size fromseveral square meters to a few square kilome-ters, and known as “bush islands”. Moststream courses are also lined with bushand/or stands of the moriche or ité palm(Mauritia flexuosa). Numerous small ponds andmarshes dot low-lying areas. The entire fringeof the savanna interdigitates with extensivehumid forest. Eden (1973) provides details onRupununi savanna geomorphology and vege-tation.

A prolonged dry season typically begins inSeptember and continues into April. Therainy season extends from late April throughAugust (83% of the annual rainfall occursduring this period; Eden 1973) and routinelyfloods much of the savanna from Junethrough early September. Not surprisingly,very little ornithological work has been con-ducted during this period. Average rainfall atDadanawa (02o49’N, 59o31’W) for 1984–1995was 1475 mm (Welle & Jansen-Jacobs 1995)and, just to the north at St. Ignatius (nearLethem 03o23’N, 59o48’W), average rainfallwas 1621 mm (Eden 1973).

With the arrival of humans, presumablythousands of years before present, the fre-quency of fire in this region increased. In1842, the Schomburgks (Roth 1922–1923)noted extensive fires set by Amerindiansthroughout the Rupununi. Fires have becomeever more frequent with the dramatic increasein cattle ranching in the late 1800s coupledwith an increase in the human populationfrom c. 5000 to 15,000 since 1950 (Hills 1976,Bureau of Statistics 1993). As is the case ingrasslands throughout the world, there isextensive burning throughout the dry season,leaving grass only a few centimeters high inmuch of the region until the rainy season. Atleast as early as the 1960s, and probably muchearlier, some areas were burned as many asthree times a year (September, December/

January, and late March/April; Hills 1976).This fire frequency continues today (D.deFreitas pers. com.; pers. observ.).

We surveyed four sites intensively during2000 (see below; Fig. 2; Appendix 1), withgeneral observations made throughout muchof this region between 17 March–22 Apriland 14 October–13 November. In addition,Finch made 10 trips to Dadanawa Ranch,which we treat as a fifth study site (Appendix1).

Karaudanawa. This camp was located along theupper reaches of the Rupununi River(02o22’N, 59o27’W), c. 4 km south of the vil-lage of Karaudanawa, and was surveyed from18 to 27 March 2000. We worked the narrowriparian corridor and the adjacent savanna.When we arrived the river was no more than afew meters across and was quite shallowexcept for a few deeper pools. However, on22 March, heavy rains fell and the river roserapidly and became impassable for three days.Several shallow ponds were present just to theeast of camp. Mauritia palms dominated alonglow-lying areas and pond edges. Primarily as aresult of burning, the grass was shortthroughout the area and there was an abruptinterface between savanna and woodland.There was very little topographical relief atthis site. From 20 to 25 12-m mist-nets wereopened from dawn until mid-morning inriparian vegetation and bush islands.

Parabara Savanna. Our camp was at the northend of this isolated savanna (02o12’N,59o22’W), just inside a belt of humid forest.This tongue of forest was 5–7 km wide wherea wagon track passed through. We surveyedsavanna and forest habitats here from 28March to 9 April 2000. The savanna had beenburned within a month prior to our arrival,and thus there was a very sharp demarcationat the savanna/forest interface. In contrast, atthe north end of the forest belt where burn-

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ing had been less intense, there was a transi-tion zone of scrubby woodland between theforest and the extensive Rupununi savanna.Most of the time we had 29 mist-netsdeployed, primarily at the south end of theforest belt and in bush islands and riparianstrips in the savanna. Finch et al. also visitedthis site on the following dates: 24–25 April1995, 13–14 April 1997, 3 November 1999,and 14 April 2001.

Wiwitau Mountain. The forest on the slopesand foot of this and adjacent granite moun-tains (02o52’N, 59o16’W) was semi-deciduous,and was surveyed from 9 to 22 April, andfrom 15 to 17 October 2000. The summitrises to 450 m. As elsewhere, most of thegrassland had been burned, leaving a sharpdemarcation between forest and savanna.From 20 to 25 mist-nets were placed in ripar-ian vegetation, bush islands, and forest at thefoot of the granite hills.

Kusad Mountain. Our camp was situated at thebase of the northeast flank of this relativelyhigh (800 m), isolated mountain (02o49’N,59o49’W), which was surveyed from 31 Octo-ber to 13 November 2000. The forest at thebase and the lower slopes was semi-decidu-ous, becoming more humid in the upper ele-vations and in narrow ravines. Most of thesurrounding savanna and some of the lowerslopes of Kusad had been extensively burned.The extent of the burning was even greater onadjacent mountains, where burns had reachedmore than half way up the slopes, destroyingmuch of the semi-deciduous forest. From 20to 25 mist-nets were placed in riparian vegeta-tion, bush islands, and in forest at the base ofthe mountain.

Dadanawa Ranch. The area which was surveyedduring 10 trips by Finch from 30 August tomid-April 1989–2001 was c. 5 km in radiusfrom the headquarters (02o49’N, 59o31’W).

Habitat is diverse, ranging from riparian for-est/scrub along the Rupununi River tosavanna with few scattered trees. There is onesemi-permanent pond at the headquarters.Like our Karaudanawa camp, there is rela-tively little topographical relief at this site. Wedid no mist-netting or collecting here, butextensive observation and tape recording overseveral years have produced a reasonablycomplete list. Mees (2000) recorded 13 spe-cies that we did not encounter at DadanawaRanch, all either water-inhabiting or forestspecies except for Long-winged Harrier (Cir-cus buffoni).

Specimens are deposited at University ofKansas Natural History Museum (KUNHM),U.S. National Museum of Natural History,Smithsonian Institution (USNM), and theUniversity of Guyana, Georgetown. Tissuesamples are at USNM. Robbins and Finchsound recordings are at Macaulay Library ofNatural Sounds, Cornell University (MLNS).Taxonomy and nomenclature mostly followHilty (2002), except for caprimulgids andhummingbirds where we follow del Hoyo etal. (1999).

RESULTS AND DISCUSSION

A total of 456 species was recorded at the fivelocalities. Of that total, 217 (referred to asuniques) were recorded at only one of the fivelocalities. The Parabara site was significantlymore diverse in total (n = 335, 73%) andunique (n = 165, 76%) species than the otherfour sites (Appendix 1). The principal factoraccounting for Parabara’s richness was thepresence of rain forest. Most of the uniqueParabara species are widespread Guiana shieldrain forest inhabitants. For example, 94% ofthe total suboscine uniques (n = 111 species)were found at Parabara. Suboscines consti-tuted 51% of the uniques among the five sites.Species totals at all sites would be increased byadditional surveys, particularly if conducted

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TABLE 1. Savanna/forest edge species sharing Peri-Atlantic pattern of distribution in selected northernSouth America savannas. Excludes species found in all six savannas. Sources: Haverschmidt & Mees(1994), Silva (1995a,1995b), Silva et al. (1997), Mees (2000), Hilty (2002), David Ascanio (pers. com.), anddata presented in this paper.

Llanos Roraima-Rupununi

Gran sabana

Sipaliwini Amapá Cerrado

Falco sparveriusBurhinus bistriatusZenaida auriculataColumbina passerinaUropelia campestrisAratinga solstitialisAratinga aureaAthene cuniculariaChordeiles pusillusHydropsalis torquataPhaethornis augustiPhaethornis rupurumiiEupetomena macrouraHeliactin bilophaPicumnus cirratusMelanerpes candidusColaptes campestrisFurnarius leucopusLepidocolaptes angustirostrisFormicivora rufaMyrmeciza longipesMyiopagis viridicataSuiriri suiririSublegatus sp.Euscarthmus rufomarginatusLophotriccus pilarisPoecilotriccus sylviaPyrocephalus rubinusXolmis cinereaXenopsaris albinuchaNeopelma pallescensAlopochelidon fucataCampylorhynchus griseusMimus saturninusPhaeothlypis flaveolaPiranga flavaEuphonia finschiNeothraupis fasciataCypsnagra hirundinaceaSporophila intermediaSporophila leucopteraSporophila bouvreuil

XXXX

XX

XX

XX

X

XXX

X

X

X

XXXX

X

XX

XX

X

X

XX

X

XXX

XX

X

XXX

X

X

XX

XX

XX

X

X

X

XX

X

XX

X

X

X

X

XX

X

XX

XXXX

X

XX

XXXXX

XX

XXX

X

X

X

X

X

XX

XX

X

X

X

XXXX

XXXXXXXX

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XXXXX

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XX

XX

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during the rainy and migratory seasons.Kusad Mountain in particular would benefitfrom visits done during the breeding season athigher elevations, as few species were vocaliz-ing there during our October–November sur-vey.

During March-April 2000, we were able todocument the breeding of about 37% of theavifauna (Appendix 1). At the three sites thatwe surveyed during March–April 2000, 147 of397 (excluding migrants, n = 9, and speciesrecorded at the Parabara site in other years)species were documented breeding based onspecimen gonad data, behavior (displays, car-rying nesting material), and location of activenests. This is a very conservative estimate ofbreeding as only 218 of the 397 species werecollected (excludes migrants and taxa repre-sented by only fluid-preserved specimenswhere gonads were not examined), and manytaxa were represented by only a single speci-men. Moreover, for some species for whichwe had reasonable sample sizes (n >10 indi-viduals), only two or three individuals were inunequivocal breeding condition (e.g., Sakes-phorus, Formicivora). Thus, larger sample sizesfor other species undoubtedly would have fur-nished additional evidence of breeding.Finally, a number of species that were not col-lected but were persistently singing, e.g., Ama-zonian Barred-Woodcreeper (Dendrocolaptescerthia), Ochre-bellied Flycatcher (Mionectes ole-agineus), Ringed Antpipit (Corythopis torquata),Bright-rumped Attila (Attila spadiceus), andBlue-black Grosbeak (Cyanocompsa cyanoides),presumably were breeding; these were not

included in our tally of documented breedingspecies.

With the exception of the ground-doves(see Species accounts), very few species weresinging during our October–November fieldwork. Nonetheless, we were able to documentbreeding by 78 of 178 (44 %) species (seeKusad locality in Appendix 1).

Below we provide details for the firstoccurrences of ten species for Guyana andthe first definite Guyana breeding recordsfor White-throated Kingbird (Tyrannus albogu-laris). The Sun Parakeet (Aratinga solstitialis)now appears extirpated in the country andthe Crestless Curassow (Crax tomentosa) hasdisappeared from much of the southernRupununi.

Intersavanna comparisons In Silva’s (1995a) biogeographic analysis ofthe cerrado avifauna, species composition iscompared across six major savanna regions ofnorthern South America that shared whatSilva termed the Peri-Atlantic pattern ofdistribution. In that analysis, he lumpedthe Sipaliwini savanna of southern Surinamewith the savannas of southeastern Venezuela[Gran Sabana; see Hilty (2002) for descriptionand distribution of this area] and contig-uous Brazil and Guyana (Roraima-Rupununi).However, with a detailed study of the avifaunaat an Amapá savanna, one of the largestsavanna islands in Brazilian Amazonia,Silva et al. (1997) documented the first occur-rence for many species in the Amapásavanna/forest edge. Their analysis of the

TABLE 1. Continuation.

Llanos Roraima-Rupununi

Gran sabana

Sipaliwini Amapá Cerrado

Sturnella magnaCarduelis cucullata

Species total

X

18

XX

26

X

8 17

X

27 31

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distribution patterns of the savanna-inhabit-ing component demonstrated that the Amapáregion’s avifauna is largely derived from thecerrado, whereas the Roraima-Rupununiavifauna is more influenced by the llanosof Colombia and Venezuela (Silva 1995b,1998).

Haverschmidt & Mees (1994) first recog-nized that the Sipaliwini savanna avifauna wascomposed mostly of cerrado species. Mees(2000) pointed out in his comparison of theGuyana Rupununi and Sipaliwini that a num-ber of species are known from the Sipaliwiniand not from the Rupununi and vice versa.Indeed, excluding water-inhabiting, strictlyriparian species, humid forest species, andmigrants, 14 species found in the Sipaliwiniand the Amapá savanna of northeastern Bra-zil (Fig. 1) are not known from the Roraima-Rupununi or the Gran Sabana savannas(Table 1). Twenty-three species are recordedfor the Roraima-Rupununi savanna and notfor the Sipaliwini, with 16 of those also unre-corded for Amapá (Table 1).

Of the species included in Table 1, not asingle species is shared by the Gran Sabanaand Sipaliwini. In contrast, the Sipaliwini avi-fauna is entirely a subset of the Amapá avi-fauna and except for the Tawny-headedSwallow [Alopochelidon (Stelgidopteryx) fucata]the Gran Sabana avifauna is a subset of theRoraima-Rupununi (Table 1). Nineteen spe-cies are known from Roraima-Rupununi, butunrecorded in the Gran Sabana [Table 1;three species known only from Santa Elenade Uairén in Venezuela (Hilty 2002) areexcluded from the Gran Sabana list].

Thus, our results from the Guyana Rupu-nuni further support Silva’s (1995b, 1998)assertion that the Roraima-Rupununi region’savifauna is most closely associated with theGran Sabana and llanos (Table 1; also see Gal-bula Species account). In contrast, the avi-fauna of the Sipaliwini savanna [considered bySilva et al. (1997) and others as part of the

larger Parú-Trombetas savanna complex ofnorthern Brazil] has closer affinities with theAmapá savanna and hence the cerrado (Table1).

Species accounts (* new for Guyana)*Broad-winged Hawk (Buteo platypterus). Anadult bird was seen circling low over a ridge atKusad Mt. on 1 November 2000 by Robbins.Although no specimen or photographicdocumentation exist for Guyana, there isan additional sight record. Finch, T. MacClen-don, and T. Thorne observed an immatureon 3 March 1994 c. 16 km west of MangoLanding, W bank Essequibo River (c.05o16’N, 59o00’W). Although not acceptedby Haverschmidt & Mees (1994), there arereliable observations for Suriname. In addi-tion to the two 1981 observations given byDonahue (1985), Finch and S. Allen observedan adult at close range at Browns BergReserve (04o53’N, 55o13’W) on 11 November1982.

Black-and-white Hawk-Eagle (Spizastur melano-leucus). Even in large expanses of lowlandand foothill forest this is a low-density spe-cies, thus we were surprised to find it in theisolated, relatively small forested hills in thesavannas. An adult was seen and heard atRaad Mountain (02o35’N, 59o51’W) on 26October 2000 by Robbins, and another adultwas present at our Kusad camp in Nov 2000.Two additional observations at the savanna/woodland interface were made by Finch in1994 near the mouth of the Ireng River (c.03o34’N, 59o52’W) and just south of Lethem.

*Merlin (Falco columbarius). We saw two atKaraudanawa in late March 2000, and twoothers at Kusad in November 2000. Althoughspecimen or photographic documentation isstill lacking for Guyana, we have encounteredthis species on no fewer than 12 occasions atboth coastal and inland localities, on dates

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ranging from 4 November to 21 April. Snyder(1966) questioned the only previous publishedreport, a seemingly very early sight record on6 August 1959. With a single exception, allVenezuela records are from north of the RíoOrinoco (Hilty 2002).

Crestless Curassow (Crax tomentosa). Mees(2000) did not find this species in the southRupununi, nor did we. This is somewhatsurprising in light of reports by Schomburgkand party (Roth 1922–1923), who mentionedthat this cracid was common in savannawoodland and riparian forest. Presumably thisdecline is the result of hunting pressure. Itstill persists at least locally in Guyana, asFinch has repeatedly heard and recorded(MLNS 78004, 87581, 106490) this cracidfrom mid-September through April in theKaranambu area as far down the RupununiRiver as the Rewa Village (03o52.1’N,58o53.6’W), and it was found in low stature,seasonally flooded forest in the Iwokramareserve (Anonymous 1999).

Ocellated Crake (Micropygia schomburgkii).This often-overlooked species was uncom-mon in the same low-lying, unburned grasswhere Polystictus was found at our Karaudan-awa camp, and in wet savanna at Parabara. Atthe latter locality at least seven individualswere heard. In addition to the Rupununirecords, this secretive crake was flushed andheard calling daily in unburned grassland withscattered shrubs at Dubulay Ranch, west bankof the Berbice River (05o40’N, 57o53’W), inlate March–early April 1996 (Robbins, C.Milensky, N. Rice, and B. Schmidt). This spe-cies is a good indicator of high quality wetgrassland and it undoubtedly was more com-mon and widespread in this region prior tothe advent of extensive, annual burning. Cav-alcanti (1988) identified this crake as a grass-land species of particular conservationconcern.

*Hudsonian Godwit (Limosa haemastica). Theonly Guyana record is of a single birdobserved and heard in flight over Musu Lake,near the Ireng River, north of Lethem on 23September 1995 by Finch et al.

*Stilt Sandpiper (Calidris himantopus). Twobirds were observed at Dadanawa Ranch on26 September 1995 by Finch. There is onlyone other record for Guyana, an observationof two birds on coastal mudflats at WindsorForest, west bank Demerara River, on 17 Sep-tember 1994 by Finch et al.

Plain-breasted Ground-Dove (Columbinaminuta). Interestingly, we failed to encounterthis species during March–April 2000, but inOctober–November 2000 it was ubiquitousand breeding throughout the south Rupununisavanna, as were the other three ground-doves and Eared Dove (Zenaida auriculata).

Sun Parakeet (Aratinga solstitialis). Discourag-ingly, we failed to find this species which isunder pressure from the pet trade. Apparentlyit was once common, at least in the northRupununi and adjacent Brazil (Roth 1922–1923, Joseph 1992), but now it is almost cer-tainly extirpated from Guyana. Rupununi birdtrappers related to us in 2000 that the specieswas once common and persisted in the northRupununi through the 1970s. Purportedly,223 individuals were exported from Guyanain 1979 (Niles 1981). However, these birdsmay well have been trapped in Brazil andbrought across the largely unpoliced border atBonfim/Lethem. As recently as 2000, weobserved relatively large numbers of parrots(Ara, Amazona, Pionus) being transportedfrom this border to Georgetown.

None of the trappers recalled havingfound the bird in the south Rupununi, nor didany of a number of long-term residents seemto know the bird. However, given that it isfound in a wide range of habitats, including

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habitat very similar to what is found through-out much of the south Rupununi, and isknown to move seasonally (Joseph 1992), wesuspect that it was once there. This species isnow of great conservation concern since,under continuing pressure from trappers, ithas disappeared from much of its formerrange.

Amazonian Pygmy-Owl (Glaucidium hardyi).This often-overlooked owl was found in thenarrow strip of humid forest at our Parabaracamp. At least 6 individuals were heard andtwo were netted (KUNHM 90816; USNM616218). The species is now known frommany Guyana localities: Kamakusa (Griscom1931), Kanuku Mts. (Parker et al. 1993,Finch), Iwokrama (Anonymous 1999), andalong the Rewa, Kuyuwini, Kassikaityu, Esse-quibo, and Kwitaro rivers in southwesternGuyana (Finch et al.).

Least Nighthawk (Chordeiles pusillus). Thispoorly known nighthawk was common inmuch of the south Rupununi from at leastOctober through April. Males collected inMarch–April had small testes, but at least twofemales taken during this period had recentlylaid eggs. Males were recorded singing anddisplaying from rocky hillside savanna inOctober (MLNS 111864). Birds foraging oversavanna during March–April evenings were inloosely associated groups numbering up to 20individuals. Although there has been very lit-tle work during the rainy season, there is aspecimen from Annai taken on 9 July 1889(American Museum of Natural History,476961) that is referable to septentrionalis, theonly subspecies documented for Guyana. So,we presume the species is a resident in thecountry.

Although Dickerman [1988; repeated byCleere (1998)] described this species’ range asincluding most of Guyana and other vastareas of inappropriate habitat in Venezuela,

Brazil, and Suriname (where it is stillunknown; Haverschmidt & Mees 1994),within Guyana this species is known onlyfrom the Rupununi region. Unless one ormore populations of this species are migra-tory, as suggested by Friedmann (1948), thedistributions that Dickerman (1988) summa-rized for the subspecies septentrionalis andesmeraldae make no sense biogeographically.The type localities are only c. 250 km apartand a specimen identified by Dickerman asbelonging to septentrionalis (Sanariapo, Amazo-nas, Venezuela) was taken within 100 km ofthe type of esmeraldae. Clearly, more material isneeded to determine if some populations aremigratory.

Streak-throated Hermit (Phaethornis rupuru-mii). Although this species was not found atany of the five sites, Finch recorded displayingmales in November and April (MLNS106313-4) at Waurepau (Wmbitun) Creek(02o20.5’N, 59o26.9’W), just south of ourKaraudanawa camp. Males sang an incessant“eesee-eesee-eesee, eesee-eesee-eesee-swur”from perches 1–1.5 m high in a dense, largelyleafless, streamside thicket within a narrowtongue of riparian woodland surrounded bysavanna. Snyder (1966) mentions that Whitelycollected the species at Annai and along theRupununi River. Meyer de Schauensee (1966)treated this taxon as a subspecies of Dusky-throated Hermit (P. squalidus), but Hinkel-mann & Schuchmann (1997) demonstrated,based on external morphological characters,that rupurumii deserves species status and ismore closely allied with Little Hermit (P.longuemareus) and Minute Hermit (P. idaliae)than with Dusky-throated.

*Versicolored Emerald (Amazilia versicolor).This species was recorded at four of our sitesat the edge of shrubby forest. Snyder (1966)did not list this species for Guyana, but Brauncollected the first specimen for the country at

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Gunn’s Strip (01o39’N, 58o37’W) in March1999 (USNM 625395). Moskovits et al. (1985)considered the species to be common in adja-cent Brazil.

Jacamars (Galbula sp.). Green-tailed Jacamar(Galbula galbula) was recorded at three of ourstudy sites (Appendix 1) and along the TakutuRiver (02o44’N, 59o59’W) and Little SandCreek, Takutu drainage (02o54’N, 59o51’W),whereas Rufous-tailed (G. ruficauda) wasrecorded only from the mouth of the IrengRiver (multiple observations by Finch et al.).Within the Rupununi, Snyder (1966) mentionsgalbula from Annai, Kanuku Mts., and theRupununi River, and Mees (2000) consideredit “moderately common and widely-distrib-uted” in tall scrub and riparian woodland inthe south Rupununi. Haffer (1974) listedspecimen records for both species in adjacentBrazil, to which may be added a report of gal-bula west of Boa Vista, along the UraricoeraRiver (Moskovits et al. 1985), and two sight-ings by Finch: galbula at Fazenda Santa Cecilia,c. 20 km southeast of Boa Vista on 19 July1992, and ruficauda near the Takutu Riverbridge on the road between Normandia andBonfim, 22 July 1992. Haffer (1974) found noevidence of hybridization where the ranges ofgalbula and ruficauda meet, and our recent spec-imens (n = 6) of galbula from the above southRupununi localities also show no signs ofhybridization.

The largely complementary ranges ofGreen-tailed and Rufous-tailed jacamars werecited as an example of competitive exclusionby Haffer (1974), although he noted that theywere sympatric in the Rio Branco drainage ofthe Roraima-Rupununi region. However, ourGuyana observations revealed no syntopy andthe above Brazilian records are consistentwith the possibility that ruficauda replaces gal-bula in the upper Rio Branco drainage. Theonly site attributed to both species is the oldcollecting locality of Forte de São Joaquim (=

Forte do Rio Branco; 03o01’N, 60o28’W; Fig.2). It may be that ruficauda occurs only to thenorth and galbula only to the south of this site.Haffer (1974) postulated that the presence ofthe nominate form of the Rufous-tailed Jaca-mar in this region was the result of pastsavanna connections with coastal Venezuelaand Guyana. Indeed, like nominate ruficauda, anumber of other avian taxa in Roraima-Rupu-nuni have disjunct populations or sister taxain northern Venezuela and/or coastal Guyanaand Suriname (Silva 1998, Mees 2000, Hilty2002, Robbins et al. 2003).

*Hoary-throated Spinetail (Poecilurus kollari).This Rio Branco drainage-restricted specieswas recorded on seven occasions, from 1993through 1996, by Finch et al., while floating c.4–5 river km along the Guyana side of theIreng River down to its confluence with theTakutu River. They encountered a maximumof 5 individuals during a float, with all birdsfound along the riverbank in dense, tangledvines that were leafless up to c. 4 m above thewater surface (MLNS 63925, 63927, 77957,78014). The species was again recorded at theconfluence when Braun collected an individ-ual on 14 November 2000 (USNM 627354).An earlier, unpublished Guyana record is thatof an unsexed individual collected along theTakutu River by D. Vesey-Fitzgerald on 17December 1932 (USNM 333145).

*Streak-headed Woodcreeper (Lepidocolaptessouleyetii). Phelps (1938) pointed out that ear-lier authors had mistakenly included Guyanawithin this species’ range, based on specimensthat were actually taken on the Venezuelanside of Mt. Roraima. Thus, the first record forGuyana was when Finch found it in riparianforest near the mouth of the Ireng River (c.03o34’N, 59o52’W), just north of Lethem on21 January 1993. At this same locality, the firstspecimens were obtained on 14 November2000 (KUNHM 91507, USNM 627537). In

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2002, this species also was found to be fairlycommon in extreme northwestern Guyana(08o15’N, 59o44’W; 08o24’N, 59o45’W) byRobbins, Milensky, and B. Benz. The Irengspecimens appear indistinguishable from theseries (n = 4) taken in northwestern Guyana.This species is known from adjacent Brazil(Sick 1993).

*Rufous-winged Antwren (Herpsilochmus rufo-marginatus). This antwren was uncommon inriparian forest canopy along the RupununiRiver (Karaudanawa camp) and the TakutuRiver (from 02o39’N, 59o58’W to 02o55’N,59o58’W) and fairly common in humid, higherelevation forest at Kusad. Snyder (1966)did not list this species for Guyana, nor arethere published records for adjacent Venezu-ela (Hilty 2002) and the Roraima state of Bra-zil (Sick 1993). However, this species does infact occur not far to the west of Guyana, asFinch recorded it along Ruta 10 at the RíoCuyuní in Bolívar, Venezuela, in 1991 and1992, and at Fazenda Santa Cecilia, about 20km southeast of Boa Vista, on 19 July 1992.We first recorded this species for Guyanain August 1998, when it was found to be fairlycommon in montane forest in the AcariMountains along the country’s southernborder.

*Rio Branco Antbird (Cercomacra carbonaria).Restricted to the Rio Branco drainage, thisspecies was not known for Guyana untilfound by Finch and T. MacClendon along thelower Ireng River on 21 January 1993. Here itoccurred in dense, low shrubby vegetationextending out over the river, and one or twoindividuals were found on five of six subse-quent visits (MLNS 63928-9, 68483, 78012-3;see Poecilurus account).

Greenish Elaenia (Myiopagis viridicata). Thisspecies was unknown in Guyana prior to Sep-tember 1989, when G. Mees found it at

Dadanawa and along the Takutu River (Mees2000). It appears to be widespread anduncommon to fairly common in semi-decidu-ous woodland, bush islands, and gallery forestthroughout much of the south Rupununi.North of the Kanuku Mountains its rangeextends at least as far as Karanambu.

Bearded Tachuri (Polystictus pectoralis). Wefound this inconspicuous tyrannid only at ourKaraudanawa camp, where it was uncommonin wet, low-lying grassland (one of the fewunburned areas). Birds were in nonbreedingcondition in March. Finch recorded this spe-cies in the southern Rupununi at MountainPoint (02o58’N, 59o39’W) and betweenDadanawa and Parabara, and in the northRupununi just north of Pirara (03o37’N,59o40’W). On 11 April 1997, birds at Moun-tain Point were in song and tape-responsive(Finch pers. observ.).

Collar & Wege (1995) listed only threeGuyana localities, but in addition to the aboverecent records, Robbins, Milensky, Rice, andSchmidt found it to be uncommon inunburned grassland with scattered scrubs atthe Dubulay Ranch in late March–early April1996. At the Dubulay site breeding apparentlyhad occurred in the past few months asimmatures with bursa of Fabricius up to 4 x 2mm were collected. Interestingly, an apparentadult female (KUMNH 88210; no bursa, skullossified, ovary mass 5 x 4 mm) had blackstreaking on the upper throat and below theeye, not unlike an adult male. We presumethat this was an old female that had acquiredmale-like characters, as other adult femalestaken in Guyana lack any black in the throatand face.

As Parker & Willis (1997) pointed out,this species has suffered considerably overmuch of its range as a result of the burning ofgrassland and its conversion to cropland. Pre-sumably it was more widespread and com-mon in the Rupununi before burning became

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a widespread and annual practice.

Yellow-olive Flatbill (Tolmomyias sulphu-rescens). We found this species to be uncom-mon to fairly common in riparian and foothillsemi-deciduous forest at several sites. Prior toour south Rupununi work and surveys byAcademy of Natural Sciences of Philadelphiapersonnel (ANSP) at Iwokrama, where thespecies was found to be uncommon in sea-sonally flooded forest (Anonymous 1999), thestatus and distribution of this species wasunknown in Guyana (Snyder 1966).

Bran-colored Flycatcher (Myiophobus fasciatus).This local species was recorded at four of oursites, where it was uncommon. Breeding wasdocumented during the March/April work(KUNHM 90774-5 – adult male; immature,bursa 3 x 1 mm). Snyder (1966) listed the spe-cies as hypothetical for Guyana, however, thefirst definite record was obtained when afemale was mist-netted on 4 July 1970, nearthe coast at Ituni, West Demerara District byJ. Dick (Dick & Barlow 1972). In addition, wemist-netted an adult male in breeding condi-tion (USNM 610189) on 8 April 1996 atDubulay Ranch. The species is known fromthe Roraima savanna in adjacent Brazil(Moskovits et al. 1985).

Variegated Flycatcher (Empidonomus varius).The northern breeding limits of this enig-matic flycatcher are poorly understood as aresult of austral migrants (nominate subspe-cies) appearing in the northern half of thecontinent from March through September(Zimmer 1937, Ridgely & Tudor 1994).Ridgely and Tudor mentioned that it is resi-dent from Venezuela south, but Hilty (2002)stated “Not yet reported breeding in Venezu-ela.” The only prior breeding record for Guy-ana is of a nest that was found on 20 April(unspecified year and locality) (Beebe et al.1917). Snyder (1966) remarked that austral

migrants “should occur” in Guyana, but vir-tually all of the localities that she listed are atinappropriate breeding sites. Thus, we suspectthat most of those records pertain to australmigrants. We are unaware of any breedingrecords for the Roraima state of Brazil. InSuriname, the resident subspecies, rufinus,apparently is found only in open savannas andhas not been recorded breeding along thecoast (Haverschmidt & Mees 1994).

We documented breeding in the southernRupununi when, on 19 March 2000, a malewas collected at Karaudanawa with testes 11 x4.5 mm (KUNHM 90782) and, the followingday, a female (USNM 625961) with an ovarymass of 8 x 5 mm with largest ovum 2 mmwas taken; based on plumage characters bothwere assignable to resident rufinus. In addition,a presumed male was recorded singing daily atdawn from the same tree at our Parabaracamp (MBR recording; MLNS 105897). Anest was found under construction and near-ing completion at the Parabara camp on 14April 2001 by Finch, J. Green, and R. Bran-son. A male collected on 9 April 1996 atDubulay Ranch (see Ocellated Crake account;KUNHM 88221) is clearly the nominate sub-species. As many as 10 individuals were seenin a day at the latter locality; we presume thatmost were austral migrants.

*Crowned Slaty Flycatcher (Griseotyrannusaurantioatrocristatus). A single bird was pho-tographed by Finch and P. Rothstein on 22April 1995, just south of the Sawariwau Riverat Mountain Point. This is the only record forGuyana, and although it is apparently a regu-lar austral migrant to Venezuela (Hilty 2002),it is still unknown from Suriname (Haver-schmidt & Mees 1994) and Sick (1993) lists norecords for Roraima savannas.

White-throated Kingbird (Tyrannus albogularis).Although Ridgely & Tudor (1994) indicatethat this species is a permanent resident

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across almost all of Guyana, southeastern-Venezuela, and the Brazilian state of Roraima,there is no concrete evidence that this king-bird, whose southern populations are migra-tory and reach northern South America,breeds in these areas (Snyder 1966, Sick 1993,Silva 1998, Hilty 2002). However, in Marchand April 2000, we documented that the spe-cies does indeed breed in at least the southernRupununi savanna. Males, with presumedmates, had enlarged testes, e.g., 10.5 x 5.5mm, 17 x 7 mm (KUNHM 90783; USNM625784, respectively), and a female takenon 20 April at Wiwitau had recently laideggs, as she had an enlarged oviduct (10 mmin diameter) and three collapsed follicles.Finch, R. Branson, and J. Green observedone carrying nest material to a morichepalm c. 10 km south of Aishalton on 13 April2001.

We did not detect this species at DubulayRanch (see Ocellated Crake account) in lateMarch–early April 1996, so the species maybe restricted as a breeder in Guyana to theRupununi savanna. Based on our Rupununidata, this species presumably breeds in savan-nas of adjacent Roraima, Brazil, and Hilty(2002) is almost certainly correct in assumingthat it breeds in the vicinity of Santa Elena deUairén, in extreme southeastern Venezuela atthe Roraima, Brazil border.

White-naped Xenopsaris (Xenopsaris albi-nucha). We found this species during Octo-ber–November 2000 in mesic, low-lying areasin the savanna. Apparently breeding hadoccurred in the past couple of months, as wecollected recently fledged young and theadults were in very worn plumage. There arethree prior Guyana records, all from theRupununi: Finch et al. studied an individual atMountain Point on 23 April 1995; one wascollected in seasonally flooded savanna nearAnnai in July 1997 (ANSP 187776); andanother was observed just north of Surama,

north Rupununi on 2 March 1998 (Anony-mous 1999). The species is known from con-tiguous Brazilian savanna (Sick 1993).

Amazonian Umbrellabird (Cephalopterus orna-tus). A single bird was observed by Finch, L.Ignacio, J. Green, and R. Branson on 14 April2001 in scrubby transitional forest borderingthe big patch of bush at the north end of theParabara savanna. The only other record forGuyana is of a specimen that was collected inthe Kanuku Mountains (Cabanis 1848), butParker et al. (1993) did not encounter it in theKanukus. Moskovits et al. (1985) recorded thespecies in adjacent Roraima, Brazil.

White-fronted Manakin (Lepidothrix serena).This species was found to be uncommon inthe narrow belt of humid forest at our Para-bara camp. This appreciably extends theknown range of the species to the west of thedistribution outlined by Prum (1994). Withthis discovery, the White-fronted Manakinand its sister species, the Orange-bellied(Tepui) Manakin (L. suavissima), which isknown from the Pakaraima Mountains ofwestern Guyana, are now documented tooccur within c. 150 km of each other.

Pale-bellied Tyrant-Manakin (Neopelma palle-scens). A totally unexpected addition to theGuyana avifauna occurred when Mees (2000)netted a male, in nonbreeding condition, on31 January 1992 in tall riparian scrub atDadanawa. Finch has found the bird at theabove locality on several occasions as well asin the north Rupununi at Karanambu. Wenetted two additional individuals (KUNHM90896; USNM 626218) at Wiwitau in March2000 that were in nonbreeding condition; nobirds were heard vocalizing nor responded totape playback during that period. Theserecords represent a range extension from thenorth bank of the Rio Amazonas near the RioTapajós area (Sick 1993).

186


Saffron Finch (Sicalis flaveola). Finch et al.observed a single individual at the Dadanawaheadquarters in November 1999 and J. Greenand R. Branson found another at MountainPoint in April 2001. Snyder (1966) mentionedthat Schomburgk and Quelch found the birdalong the coast in the 1800s, and prior to theFinch record, P. Peberdy was the last to recordthe species for Guyana when in 1938 hereported it from the Marudi Mountains, justto the southeast of the Rupununi. It isunknown from adjacent Brazil (Sick 1993)and the Gran Sabana (Hilty 2002).

ACKNOWLEDGMENTS

The American Bird Conservancy, AmericanFederation of Aviculturists, Bushnell SportsOptics, Conservation International, andNational Aviary supported in part our Octo-ber and November 2000 fieldwork. TheNational Biodiversity Committee of the Envi-ronmental Protection Agency, the Ministry ofAmerindian Affairs, and the University ofGuyana provided permission and help for ourwork. We thank our field colleagues BrianSchmidt, Chris Milensky, Brian Barber,Nathan Rice, Brian O’Shea, and WiltshireHinds. Milensky (USNM), Rice (ANSP), PaulSweet, and Shannon Kenney (AMNH) pro-vided specimen information. Duane, Sandy,and Justin deFreitas, Andy Narain, AshleyHolland, and Claudius Perry helpedimmensely with logistics. Tom Hollowell,Carol Kellof, Thomas Schulenberg, and Gra-ham Watkins kindly supplied several Rupu-nuni publications. Hollowell also producedboth maps. Greg Budney and Martha Fischerprovided catalog data for recordings depos-ited at the Macaulay Library of NaturalSounds, Cornell University (MLNS). JürgenHaffer, José Maria Cardoso da Silva, andDavid Ascanio made valuable comments thatimproved the manuscript. Adolfo Navarro-Sigüenza and Town Peterson translated the

abstract. This is number 78 in the Smithso-nian’s Biological Diversity of the Guianas Pro-gram publication series.

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APPENDIX 1. Relative abundance and breeding status of Rupununi birds of five sites. Relative abun-dance: C = common, more than 20 individuals/day; F = fairly common, 5–20 individuals/day; U =uncommon, present in small numbers (< 5 individuals/day) but not encountered daily, even in prime habi-tat; S = scarce, only occasionally encountered in small numbers; X = Single record. Asterisk indicatesbreeding during March/April; # indicates breeding during October/November period.

Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusTinamidae

Tinamus majorCrypturellus cinereusCrypturellus souiCrypturellus undulatusCrypturellus erythropusC. erythropus/undulatusCrypturellus variegatus

PhalacrocoracidaePhalacrocorax brasilianus

AnhingidaeAnhinga anhinga

ArdeidaeBotaurus pinnatusIxobrychus exilisArdea cocoiArdea albaEgretta thula

SS

X

XS

FUU

F

U

SXS

X

U

S

S

F

U

S

FFU

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APPENDIX 1. Continuation.

Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusEgretta caeruleaBubulcus ibisButorides striatusPilherodius pileatusNycticorax nycticorax

ThreskiornithidaeCercibis oxycercaTheristicus caudatusMesembrinibis cayennensis

CiconiidaeJabiru mycteriaMycteria americana

CathartidaeCoragyps atratusCathartes auraCathartes burrovianusCathartes melambrotusSarcoramphus papa

AnatidaeDendrocygna viduataCairina moschataAmazonetta brasiliensisOxyura dominica

AccipitridaePandion haliaetusLeptodon cayanensisElanoides forficatusGampsonyx swainsoniiElanus leucurusIctinia plumbeaBusarellus nigricollisAccipiter bicolorGeranospiza caerulescensLeucopternis albicollisButeogallus urubitingaButeogallus meridionalisButeo nitidusButeo magnirostrisButeo platypterusButeo brachyurusButeo albicaudatusButeo albonotatusSpizastur melanoleucusSpizaetus tyrannus

FalconidaeMicrastur ruficollis

XXX

S

FFU

S

S

X

XXX

US

X

XUXU

S

X

UF

UU

X

F

U

X

U

F

U

XS

U

X

U

UU

CFF

F

U

SS

SS

SUSU

SFU

F

X

UFU

S

U

SU

SSFSFX

UXX

S

SCUXU

UFF

F

CFF

U

UFU

U

XUXUU

UF

F

U

*

#

*

*

#

**

*

190



Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusMicrastur gilvicollisMicrastur semitorquatusDaptrius aterIbycter americanusCaracara cheriwayMilvago chimachimaHerpetotheres cachinnansFalco sparveriusFalco columbariusFalco femoralisFalco rufigularisFalco deiroleucusFalco peregrinus

CracidaeOrtalis motmotPipile cumanensisCrax alector

OdontophoridaeColinus cristatusOdontophorus gujanensis

RallidaeMicropygia schomburgkiiAnurolimnas viridisAramides cajaneaPorzana albicollisPorphyrio martinicaPorphyrio flavirostris

EurypygidaeEurypyga helias

AramidaeAramus guarauna

PsophidaePsophia crepitans

BurhinidaeBurhinus bistriatus

CharadriidaeHoploxypterus cayanusVanellus chilensisPluvialis dominicaCharadrius collaris

JacanidaeJacana jacana

ScolopacidaeTringa melanoleucaTringa flavipesTringa solitaria

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191

ROBBINS ET AL.


Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusActitis maculariaBartramia longicaudaCalidris minutillaCalidris fuscicollisCalidris melanotosCalidris himantopusGallinago paraguaiaeGallinago undulata

ColumbidaeColumba speciosaColumba cayennensisColumba plumbeaColumba subvinaceaZenaida auriculataColumbina passerinaColumbina minutaColumbina talpacotiClaravis pretiosaLeptotila verreauxiLeptotila rufaxillaGeotrygon montana

PsittacidaeAratinga pertinaxAra chloropteraAra macaoAra araraunaOrthopsittaca manilataDiopsittaca nobilisBrotogeris chrysopterusTouit sp.Pionites melanocephalaPionopsitta caicaPionus menstruusPionus fuscusAmazona ochrocephalaAmazona amazonicaDeroptyus accipitrinus

CuculidaePiaya cayanaTapera naeviaCrotophaga ani

StrigidaeOtus cholibaOtus watsoniiLophostrix cristataPulsatrix perspicillata

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192



Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusBubo virginianusGlaucidium hardyiGlaucidium brasilianumAthene cuniculariaStrix sp.Asio stygius

CaprimulgidaeChordeiles pusillusChordeiles acutipennisPodager nacundaNyctidromus albicollisCaprimulgus cayennensis

NyctibiidaeNyctibius grandisNyctibius griseus

ApodidaeStreptoprogne zonarisChaetura brachyuraChaetura spinicaudaChaetura cinereiventrisPanyptila cayennensisTachornis squamata

TrochilidaePhaethornis superciliosusPhaethornis bourcieriPhaethornis augustiPhaethornis ruberFlorisuga mellivoraAnthracothorax nigricollisChrysolampis mosquitusLophornis ornatusChlorestes notatusChlorostilbon mellisugusThalurania furcataHylocharis sapphirinaHylocharis cyanusPolytmus guainumbiPolytmus theresiaeAmazilia versicolorAmazilia fimbriataAmazilia cupreicaudaHeliothryx auritaHeliomaster longirostrisCalliphlox amethystina

TrogonidaeTrogon viridis

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193

ROBBINS ET AL.


Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusTrogon violaceusTrogon rufusTrogon melanurus

MomotidaeMomotus momota

AlcedinidaeCeryle torquataChloroceryle amazonaChloroceryle americanaChloroceryle inda

BucconidaeCapito nigerNotharchus macrorhynchosNotharchus tectusBucco tamatiaBucco capensisMalacoptila fuscaMonasa atraChelidoptera tenebrosa

GalbulidaeBrachygalba lugubrisGalbula albirostrisGalbula galbulaGalbula leucogastraGalbula deaJacamerops aureus

RamphastidaePteroglossus viridisPteroglossus aracariRamphastos vitellinusRamphastos tucanus

PicidaePicumnus cirratusDryocopus lineatusMelanerpes cruentatusVeniliornis passerinusVeniliornis cassiniPiculus flavigulaPiculus chrysochlorosCeleus elegansCeleus undatusCeleus flavusCeleus torquatusCampephilus rubricollisCampephilus melanoleucos

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194



Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusFurnariidae

Furnarius leucopusSynallaxis albescensCerthiaxis cinnamomeaPhilydor pyrrhodesPhilydor erythrocercusAutomolus ochrolaemusAutomolus infuscatusAutomolus rubiginosusAutomolus rufipileatusXenops minutusSclerurus rufigularis

DendrocolaptidaeDendrocincla fuliginosaGlyphorynchus spirurusDendrocolaptes certhiaDendrocolaptes picumnusXiphorhynchus picusXiphorhynchus pardalotusXiphorhynchus guttatusLepidocolaptes albolineatusCampylorhamphus procurvoides

ThamnophilidaeCymbilaimus lineatusFrederickena viridisTaraba majorSakesphorus canadensisThamnophilus doliatusThamnophilus murinusThamnophilus punctatusThamnophilus amazonicusThamnomanes ardesiacusThamnomanes caesiusMyrmotherula brachyuraMyrmotherula guttataMyrmotherula gutturalisMyrmotherula axillarisMyrmotherula longipennisMyrmotherula menetriesiiHerpsilochmus sticturusHerpsilochmus stictocephalusHerpsilochmus rufomarginatusFormicivora griseaTerenura spodioptilaCercomacra cinerascensCercomacra tyrannina

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195

ROBBINS ET AL.


Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusCercomacra laetaMyrmoborus leucophrysHypocnemis cantatorPercnostola rufifronsMyrmeciza ferrugineaHylophylax naeviaHylophylax poecilinotaMyrmornis torquataPithys albifronsGymnopithys rufigula

FormicariidaeFormicarius colmaFormicarius analisHylopezus maculariusMyrmothera campanisona

TyrannidaeOrnithion inermeCamptostoma obsoletumPhaeomyias murinaCapsiempis flaveolaTyrannulus elatusMyiopagis viridicataMyiopagis gaimardiiElaenia flavogasterElaenia cristataElaenia ruficepsElaenia chiriquensisInezia caudataMionectes oleagineusPolystictus pectoralisSublegatus sp.Zimmerius gracilipesMyiornis ecaudatusLophotriccus vitiosusLophotriccus galeatusLophotriccus pilarisHemitriccus josephinaePoecilotriccus sylviaTodirostrum cinereumTodirostrum pictumCorythopis torquataRamphotrigon ruficaudaRhynchocyclus olivaceusTolmomyias sulphurescensTolmomyias assimilis

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196



Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusTolmomyias poliocephalusTolmomyias flaviventrisPlatyrinchus saturatusOnychorhynchus coronatusMyiobius barbatusMyiophobus fasciatusHirundinea ferrugineaLathrotriccus euleriContopus cinereusPyrocephalus rubinusArundinicola leucocephalaColonia colonusAttila cinnamomeusAttila spadiceusRhytipterna simplexRhytipterna immundaSirystes sibilatorMyiarchus tuberculiferMyiarchus swainsoniMyiarchus feroxMyiarchus tyrannulusPhilohydor lictorPitangus sulphuratusMegarynchus pitanguaMyiozetetes cayanensisConopias parvaMyiodynastes maculatusLegatus leucophaiusEmpidonomus variusTyrannopsis sulphureaTyrannus albogularisTyrannus melancholicusTyrannus savanaSchiffornis turdinusPiprites chlorisLipaugus vociferansLaniocera hypopyrraXenopsaris albinuchaPachyramphus polychopterusPachyramphus marginatusPachyramphus rufusPachyramphus minorTityra cayanaTityra inquisitor

CotingidaeCotinga cotinga

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197

ROBBINS ET AL.


Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusCotinga cayanaXipholena puniceaGymnoderus foetidusHaematoderus militarisQuerula purpurataCephalopterus ornatusPerissocephalus tricolorProcnias alba

PipridaeXenopipo atronitensManacus manacusChiroxiphia pareolaPipra pipraPipra erythrocephalaLepidothrix serenaTyranneutes virescensNeopelma pallescens

VireonidaeVireo olivaceus Hylophilus thoracicusHylophilus pectoralisHylophilus muscicapinusVireolanius leucotisCyclarhis gujanensis

CorvidaeCyanocorax cayanus

HirundinidaeProgne chalybeaProgne taperaTachycineta albiventerStelgidopteryx ruficollisRiparia ripariaHirundo rustica

TroglodytidaeCampylorhynchus griseusThryothorus corayaThryothorus leucotisTroglodytes musculusHenicorhina leucostictaMicrocerculus bamblaCyphorhinus arada

SylviidaeMicrobates collarisRamphocaenus melanurusPolioptila plumbea

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198

ROBBINS ET AL.


Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusTurdidae

Catharus fuscescensCatharus minimusTurdus leucomelasTurdus nudigenisTurdus albicollis

MimidaeMimus gilvus

MotacillidaeAnthus lutescens

ParulidaeParula pitiayumiDendroica petechiaGeothlypis aequinoctialisBasileuterus culicivorusPhaeothlypis flaveolaPhaeothlypis rivularisGranatellus pelzelni

CoerebidaeCoereba flaveola

ThraupidaeConirostrum speciosumSchistochlamys melanopisLamprospiza melanoleucaHemithraupis guiraNemosia pileataLanio fulvusTachyphonus cristatusTachyphonus surinamusTachyphonus rufusTachyphonus phoeniciusPiranga flavaPiranga rubraRamphocelus carboThraupis episcopusThraupis palmarumCyanicterus cyanicterusEuphonia chloroticaEuphonia finschiEuphonia violaceaEuphonia chrysopastaTangara chilensisTangara punctataTangara cayanaTangara veliaDacnis lineata

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199


Karaudanawa Parabara Wiwitau Kusad Dadanawa Breeding statusDacnis cayanaChlorophanes spizaCyanerpes nitidusCyanerpes caeruleusCyanerpes cyaneus

EmberizidaeVolatinia jacarinaSporophila intermediaSporophila plumbeaSporophila nigricollisSporophila minutaOryzoborus crassirostrisOryzoborus angolensisDolospingus fringilloidesSicalis flaveolaSicalis luteolaEmberizoides herbicolaParoaria gularisArremon taciturnusAmmodramus humeralis

CardinalidaePitylus grossusSaltator coerulescensCaryothraustes canadensisCyanocompsa cyanoides

IcteridaeSturnella militarisSturnella magnaMolothrus bonariensisMolothrus oryzivorusIcterus chrysocephalusIcterus cayanensisIcterus nigrogularisCacicus celaCacicus haemorrhousPsarocolius decumanusPsarocolius viridis

FringillidaeCarduelis cucullata

Total speciesUniques/site

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FUFXF

U

F

F

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SCU

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1886

UCUCC

C

U

FX

F

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XCSU

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335165

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F

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UC

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1497

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C

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17813

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SFXU

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18526

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200

avifauna of the guyana southern rupununi...

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