Handbook birds world volume 16 by Lynx Edicions

Volume 16 Tanagers to New World Blackbirds

001-Portadelles-HBW-16.p65

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ssp cucullata ssp saturata ssp montana

ssp eximia

86 85 84 83 ssp chloronota

ssp lacrymosus

ssp palpebrosus

87 ssp erythrotus ssp intensus

ssp igniventris ssp pallididorsalis

ssp somptuosus

PLATE 9

89 ssp victorini

ssp baezae ssp cyanopterus

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inches

Family THRAUPIDAE (TANAGERS)

201

SPECIES ACCOUNTS

PLATE 9

Genus WETMORETHRAUPIS Lowery & O’Neill, 1964

81. Orange-throated Tanager Wetmorethraupis sterrhopteron French: Tangara à gorge orangée Spanish: Tangara Golinaranja German: Veilchenschultertangare Other common names: Wetmore’s Tanager Taxonomy. Wetmorethraupis sterrhopteron Lowery and O’Neill, 1964, Chávez Valdivia, Amazonas, Peru. Molecular studies place present species as sister to Bangsia; differs from other genera in family in stiff, bristly throat feathers, although this condition is approached in Spindalis. Monotypic. Distribution. Extreme S Ecuador (SE Zamora-Chinchipe) and adjacent N Peru (NC Amazonas). Descriptive notes. 17 cm; 43·6–56 g. Strikingly plumaged, thick-billed tanager with relatively long wing, but short tail and tarsus; feathers of malar region, lower chin and throat stiff, almost bristly, and with waxy appearance. Has top and side of head, lores, chin, side of neck and upperparts, including uppertail-coverts, velvety black, tail matt black; bend of wing greyish-blue, marginal coverts, lesser upperwing-coverts and lower half of median coverts deep blue, upper half of medians black, edged and tipped same blue colour, greater coverts blackish, broadly edged and tipped light violet-blue; alula and primary coverts black, narrowly edged blue; primaries black, secondaries and tertials black, edged pale violet-blue; throat and upper chest deep cadmium-orange, feathers of chest with burnished quality, rest of underparts buff-yellow, thigh black; underwing-coverts blackish-slate, a few feathers edged blue; iris dark brown; bill and legs black. Sexes similar. Juvenile apparently undescribed. VOICE. Dawn song, in Peru, a thick, heavy “jasuu, jasuu, jasuu...”, up to a half-dozen notes, repeated over and over with scarcely a pause; also (possibly also day song), from high perch, a much higher-pitched, squeaky, slurred series of 3-note phrases, e.g. “we-tsí-tsoo, we-tsí-tsoo”, described also as “inchúé-tooch”, first note high (often inaudible at distance), second syllable loudest. Call a soft, penetrating “seet”. Habitat. Mainly mature humid and wet pre-montane forest in relatively gentle foothills, and terra firme forest in adjacent lowlands; reported also in disturbed mature forest. Recorded at c. 450– 800 m in Peru; 600–1000 m in Ecuador. Food and Feeding. Fruit and insects. One stomach contained seeds, fruit pulp and a beetle (Coleoptera). Found in pairs and in small groups of up to about six individuals, foraging in middle storey to canopy; sometimes with mixed-species flocks, but more often away from them. Fairly conspicuous. Scans leaves, and searches mosses and small bromeliads as it hops along branches; has been seen while foraging in Cecropia trees, and hovering, presumably to take fruit, at tips of branches. Apparently ranges over large areas. Breeding. No information. Movements. Apparently resident. Status and Conservation. VULNERABLE. Restricted-range species: present in Andean Ridgetop Forests EBA. Notably local; fairly common in suitable habitat at a few sites. First described in 1964 in Peru, and more recently (1990) found in Ecuador. This species was first brought to the attention of scientists by indigenous hunters living in vicinity of type locality. In extreme S Ecuador, recorded above R Nangaritza on W slope of S Cordillera del Cóndor, in SE Zamora-Chinchipe. In adjacent Peru, recorded in NC Amazonas S to Urakusa on lower E slopes and foothills above R Cenepa, and from regions S of R Marañón opposite mouth of R Cenepa; reported by native Aguaruna tribe as occurring also at headwaters of R Nieva (also in Amazonas), a locality some distance SE of the species’ known range. In S Ecuador, seen regularly on Shaur indigenous land (near village of Shaime, on R Nangaritza) in Cordillera del Cóndor. Widespread forest destruction has taken place within its very small known global range in S Ecuador and N Peru, and none of the land within its range is protected as yet. The loss of some forest has occurred within the last 20 years (since 1990) within its tiny range in Ecuador, where recent authors regard the species as “Endangered”; it appears that it is in decline in all areas. Further field surveys required in order to determine the species’ precise distribution. Bibliography. Anon. (2010a), Begazo et al. (2001), Butchart & Stattersfield (2004), Capper & Pereira (2007), Clements & Shany (2001), Isler & Isler (1999), Lowery & O’Neill (1964), Lysinger et al. (2005), Marín et al. (1992), O’Neill (1969), van Oosten et al. (2007), Remsen et al. (2010), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Schulenberg et al. (2007), Sedano & Burns (2010), Stattersfield & Capper (2000).

Birds from Venezuela proposed as a separate race, venezuelana, but validity of this requires confirmation. Five subspecies currently recognized. Subspecies and Distribution. B. m. gigas (Bonaparte, 1851) – Venezuela (Sierra de Perijá, and Andes of S Táchira) and adjacent E Andes of Colombia (S to Cundinamarca). B. m. cucullata (Jardine & Selby, 1842) – W & C Andes of Colombia S to Ecuador. B. m. cyanonota Berlepsch & Stolzmann, 1896 – Andes of N & C Peru (Amazonas S to Junín). B. m. saturata Berlepsch & Stolzmann, 1906 – Andes of SE Peru (Cuzco and Puno). B. m. montana (d’Orbigny & Lafresnaye, 1837) – Andes of N Bolivia (La Paz, Cochabamba and W Santa Cruz). Descriptive notes. 21 cm, 69–91 g (nominate); 23 cm, 72–116 g (other races). Largest mountain-tanager; distinctive rotund shape and conspicuous red eyes, short and thick bill. Nominate race has head and throat black, small amount of black extending down onto side of chest and breast (often concealed), hindcrown and nape (nuchal collar) shining pale milky blue, contrasting sharply with dark shining blue back and rest of upperparts; lesser upperwing-coverts blue, median and greater coverts black, edged blue, primary coverts black, flight-feathers blackish, inner primaries and all of secondaries edged blue, tertials black, broadly edged blue; tail feathers black, edged blue; breast and lower underparts goldenyellow, narrow blue-and-black band from flank extending onto thigh (like dark garter); iris bright red; bill blackish above and below, basal half of lower mandible often paler bluish-grey; legs blackish. Sexes similar. Immature is paler and duller, especially below, than adult. Races differ in size, nominate smallest, others lack obvious pale nuchal collar, have base of lower mandible pale blue-grey (not pinkish), and differ slightly in tone of upperparts: cyanonota is closest to nominate, but larger; cucullata is shining windsor-blue above, paler than nominate; saturata is shining violaceous blue above; gigas is dull purplish-blue above. VOICE. During flight display, presumed male sings rapid, exuberant “chip’ut-chip’ut-chip-ut...” as he flies out high in large 100-m circle above forest, before diving back into forest. Apparent dawn song a single “weeck!” or “tooweeck” repeated monotonously over and over at rate of about once per second. Also gives repeated “tsee-tsee-tseét?”. Group-members, sometimes stimulated by extraneous noises, simultaneously give loud bursts of squealing notes; also, during foraging, “weeck” and “too” notes, or “tooweeck” phrases, often rapidly; also weak “ti” notes singly or in rapid series. Habitat. Mainly humid and wet montane forest and forest borders; also tall second-growth woodland, scattered trees in clearings adjacent to forest, and stunted or elfin forest and scrub woodland near tree-line. Mostly at 1900–3500 m; 1800–3000 m in Venezuela, and once (possibly a vagrant) as low as 1100 m on Pacific slope of Colombia. Food and Feeding. Arthropods, also many small berries and fruit. Of 35 stomachs examined, 34 contained only vegetable matter and one contained both animal and vegetable matter; items included fruit pulp, berries, seeds, including grass seeds (Poaceae), leaves and beetles (Coleoptera). Social, often seen in conspicuous groups of 3–10 individuals, occasionally more, that roam over large areas, and sometimes rather confiding. Flocks move along steep forested slopes or canyons, often crossing considerable gaps with each flight, stragglers hurrying along to keep up with fastermoving members. Regularly occurs in loosely associated mixed flocks with Southern Mountain (Cacicus chrysonotus) and Northern Mountain Caciques (Cacicus leucoramphus) and jays (Corvidae), when forages rapidly, but only infrequently with mixed flocks of smaller species. Away from mixed-flock associations, forages in slower methodical manner on bare or mossy branches c. 25 mm or more in diameter. Forages mainly in middle and upper strata of trees, occasionally lower. Hops up through outer-canopy limbs, leans down to inspect sides of branches, especially to examine lichens on branch surfaces; also noted as taking prey from foliage, and may examine flowers, seedpods and even dead-leaf clusters. Breeding. No information. Movements. Resident. A few scattered records, probably of wandering or vagrant birds, at elevations lower than usual. Status and Conservation. Not globally threatened. Fairly common throughout its wide latitudinal distribution in Andes. Generally occurs at rather low density, perhaps because groups wander over large areas of forest. Deforestation has fragmented this species’ range in many areas, but it occurs in many protected areas, from Venezuelan Andes S to Bolivia; among them are Guaramacal and Sierra Nevada National Parks (Venezuela), Chingaza and Puracé National Parks and Tambito Nature Reserve (Colombia), Sangay and Las Cajas National Parks (Ecuador), Tingo Maria and Manu National Parks (Peru) and Isiboro-Sécure and Amboró National Parks (Bolivia). This species is found also in many areas of unprotected montane forest in Andes. Bibliography. Álvarez-Rebolledo & Córdoba-Córdoba (2002), Aveledo Hostos & Pérez Chinchilla (1989), Burns (1997a), Burns & Naoki (2004), Burns et al. (2002, 2003), Donegan & Dávalos (1999), Dunning (1982), Fjeldså & Krabbe (1990), Hilty (2003), Hilty & Brown (1986), Isler & Isler (1999), Jacobs & Walker (1999), Krabbe et al. (2001), Remsen & Parker (1984), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman, Donegan & Caro (2008), Salaman, Donegan & Cuervo (1999), Schulenberg et al. (2007), Sedano & Burns (2010), Walker (2001), Weske (1972), Yuri & Mindell (2002), Zimmer (1944).

83. Black-chested Mountain-tanager

Genus BUTHRAUPIS

Cabanis, 1851

French: Tangara à poitrine noire Spanish: Tangara Dorsiverde German: Schwarzbrust-Bergtangare Other common names: Blue-rumped Mountain-tanager

82. Hooded Mountain-tanager Buthraupis montana French: Tangara montagnard

German: Blaurücken-Bergtangare

Buthraupis eximia

Spanish: Tangara Montana

Taxonomy. A.[glaia] montana d’Orbigny and Lafresnaye, 1837, Yungas, Bolivia.

Taxonomy. Tanagra eximia Boissonneau, 1840, Santa Fé de Bogotá, Colombia. May form a superspecies with B. aureodorsalis. Four subspecies recognized. Subspecies and Distribution. B. e. eximia (Boissonneau, 1840) – Andes of SW Venezuela (S Táchira) and adjacent Colombia (Norte de Santander and Cundinamarca).

On following pages: 84. Golden-backed Mountain-tanager (Buthraupis aureodorsalis); 85. Masked Mountain-tanager (Buthraupis wetmorei); 86. Santa Marta Mountain-tanager (Anisognathus melanogenys); 87. Lacrimose Mountain-tanager (Anisognathus lacrymosus); 88. Scarlet-bellied Mountain-tanager (Anisognathus igniventris); 89. Blue-winged Mountain-tanager (Anisognathus somptuosus); 90. Bolivian Mountain-tanager (Anisognathus flavinucha); 91. Black-chinned Mountain-tanager (Anisognathus notabilis).

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Family THRAUPIDAE (TANAGERS)

PLATE 9

SPECIES ACCOUNTS

B. e. zimmeri R. T. Moore, 1934 – Colombia at N end of W Andes (páramos of Frontino and Paramillo) and in C Andes (Caldas–Antioquia border S to Tolima). B. e. chloronota P. L. Sclater, 1855 – S Colombia on both slopes of Andes (from Huila and Cauca) S to NW Ecuador. B. e. cyanocalyptra R. T. Moore, 1934 – Andes of S Ecuador S to N Peru (S to E Piura and N Cajamarca, mostly N of R Marañón). Descriptive notes. 18–22 cm; 50–70 g. Darkplumaged, quiet tanager with short, thick bill, usually a few stiff bristles on chin. Nominate race has crown and nape silky cornflower-blue, mantle and back moss-green, rump and long uppertail-coverts silky blue (like crown); lesser and median upperwing-coverts blue (often partly concealed), greater coverts leaf-green with dark bases, flight-feathers black, secondaries edged green, tertials mostly dull green with dusky inner webs; tail black; lores, ocular area, and side of head to throat and chest black, rest of underparts golden-yellow, sometimes with sides narrowly dark mossy green (usually hidden); undertail-coverts duller; iris dark reddish-brown; bill black; legs dark grey. Sexes similar. Juvenile apparently undescribed. Race zimmeri differs from nominate in having rump mixed blue and dark green; cyanocalyptra differs in having rump and uppertail-coverts all dark moss-green, like back, and central throat with small concealed white patch; chloronota is larger than others, has duller blue head, black of mask less sharply separated from adjoining blue, rump and tail-coverts all dark green (like previous), lacks bristles on chin. VOICE. Dawn song a simple series of alternating notes, the first lower, e.g. “tchew syéeuw, tchew syéeuw, tchew syéeuw”; also a more complex song lasting up to 30 seconds, e.g. “tititi-turri-tititi-tee-ter-turry...”; neither song heard frequently. Variety of soft “seep” notes, “chip” notes and forced “piz-weet” and soft “weeur”, etc. during foraging. Habitat. Wet mossy montane forest, along forest borders and in elfin woodland near tree-line. Recorded at 2000–3800 m, mostly c. 2750–3400 in Colombia and Ecuador, and mainly 2850–3300 m in N Peru. Largely replaces B. montana at higher elevations. Food and Feeding. Mostly fruit; also insects. In Colombia recorded as taking fruit from Ericaceae, Miconia (Melastomataceae), and Hedyosmum (Chloranthaceae). Seven stomachs all contained vegetable matter, including seeds and fruit pulp. Found singly, in pairs and sometimes in small groups of 3–6 individuals, most often alone, but sometimes with mixed-species flocks. Quiet; moves rather heavily and methodically along mossy branches, mostly within dense foliage of trees and shrubs, and sometimes sits quietly, without moving, for several minutes in crown of small to large tree. In general much less conspicuous than B. montana. Searches for insects in moss and other canopy epiphytes. Breeding. No information. Movements. Presumed resident; little information. Status and Conservation. Not globally threatened. Generally uncommon to rare. Uncommon in Venezuela. Rare to uncommon and somewhat local in Ecuador and Colombia, where most numerous at higher elevations; often found in elfin woodland at tree-line. In Peru, race cyanocalyptra confined to a narrow corridor of upper montane forest in extreme N, generally N of R Marañón, but two records S of Marañón Valley. Occurs at rather low density, and deforestation has fragmented this species’ range in many areas; in Peru, deforestation and human settlement have been expanding since c. 1980. Present in numerous protected areas, among them Chingaza and Puracé National Parks (Colombia) and Cayambe, Sangay, Las Cajas and Podocarpus National Parks (Ecuador). Bibliography. Álvarez-Rebolledo et al. (2007), Anon. (2000), Barnes et al. (1997), Botero et al. (2005), Fjeldså & Krabbe (1990), Hilty (2003), Hilty & Brown (1986), Isler & Isler (1999), Krabbe, Flórez et al. (2006), Krabbe, Moore et al. (2001), Meyer de Schauensee (1964, 1966, 1970a), Moore (1934a), Olivares (1969), Parker et al. (1985), Remsen et al. (2010), Restall et al. (2006), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Schulenberg et al. (2007), Sedano & Burns (2010), Sibley & Monroe (1990).

84. Golden-backed Mountain-tanager Buthraupis aureodorsalis French: Tangara à dos d’or

German: Goldrücken-Bergtangare

Spanish: Tangara Dorsidorada

Taxonomy. Buthraupis aureodorsalis Blake and Hocking, 1974, Quilluacocha, c. 11,500 feet [c. 3500 m], Huánuco (Acomayo), Peru. May form a superspecies with B. eximia. Monotypic. Distribution. E slope of Andes in NC Peru (San Martín, La Libertad and Huánuco). Descriptive notes. 23 cm; 75–94 g. Large, spectacular but hard-to-find tanager; bill short and thick. Has crown dull blue, rest of head down to upper mantle and mid-breast black; rest of mantle, scapulars, rump and uppertail-coverts bright orange-yellow with sparse and rather obscure dusky and brownish markings; lesser upperwing-coverts blue, like crown (but often hidden), median and greater coverts, flightfeathers, tertials and tail deep black; lower breast, belly, sides and flanks bright orange-yellow (like back), belly and sides also with sprinkling of arrow-like bright chestnut spots and streaks (markings decreasing in size rearwards), undertail-coverts and lower belly chestnut, thigh commonly yellow at front but chestnut behind or on inside; axillaries bright yellow, undersurface of wing black, except for small patch of yellow on outer edge of bend of wing; iris dark brown; bill blackish above, paler below; legs dark grey. Sexes similar. Immature is similar to adult, but duller and with little blue on lesser wing-coverts. VOICE. Song a series of wheedles, squeals and churrs; typically, repeats one note 4–6 times and then switches to another in rather staccato series given hurriedly and rhythmically. Variety of sharp, metallic and sometimes squeaky notes, including “chit’, “wheet” and “heet”, singly or repeated, and thin chittering “sweet” or “sweee” notes, all of them surprisingly weak for so relatively large a bird, and frequently given by foraging groups, sometimes for lengthy periods of time; sharp “steet-steet” in flight. Habitat. Elfin forest at tree-line; especially in large islands and patches of Escallonia and Clusia woodland surrounded by grass at tree-line, and frequently along ridges enveloped in clouds. Occasionally in scattered low trees and scrub in more open areas near elfin forest. At 3150–3500 m.

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Food and Feeding. Fruits and insects. Of seven stomachs examined, five contained only vegetable matter and two contained both vegetable and animal matter; contents included fruit pulp, seeds, and a beetle (Coleoptera) 8 mm long. Usually seen in inconspicuous pairs, less often in small parties of 3–5 (occasionally up to 7) individuals, which forage alone or sometimes join small mixedspecies flocks. Regularly flies off some distance in undulating flight to new patch of woodland; typically, first bird calls as it takes flight, is quickly followed by others in rapid succession, and, once they arrive, they rest or forage. Forages from ground to treetops, mostly at middle heights in small trees and shrubs, often c. 3 m up. Moves quietly and slowly, hanging or leaning downwards to peck items from moss and lichens on branches, or occasionally from leaves. Most prey items believed to be taken from moss. Breeding. Brooding females in Sept, and juveniles and immatures found in Jul, Oct and Nov. No other information. Movements. Presumed resident; no evidence of any movements. Status and Conservation. ENDANGERED. Restricted-range species: present in North-east Peruvian Cordilleras EBA. Uncommon to locally fairly common. Estimated global population very small, and declining as a result of loss of habitat, especially of tree-line Polylepis and other trees and shrubs of elfin woodland. Currently known from five sites within restricted area of Cordillera Central in Peruvian departments of San Martín, La Libertad and Huánuco; may occur in other, unexplored regions nearby, but to date has not been found farther N (in Cordillera de Colán). Occurs at low density in Río Abiseo National Park (San Martín), where noted only three times in 27 days of field survey; possibly easiest to find in Carpish Ridge region at Quilluacocha and Bosque Unchog, in Huánuco. Continued destruction of tree-line woodlands represents a major threat to this species; spread of fires from adjacent grassland also a cause of serious concern. Bibliography. Anon. (2010a), Butchart & Stattersfield (2004), Clements & Shany (2001), Fjeldså & Krabbe (1990), Isler & Isler (1999), Remsen et al. (2010), Ridgely & Tudor (1989, 2009), Schulenberg et al. (2007), Sedano & Burns (2010), Sibley & Monroe (1990), Stattersfield & Capper (2000).

85. Masked Mountain-tanager Buthraupis wetmorei French: Tangara de Wetmore

German: Wetmorebergtangare

Spanish: Tangara de Wetmore

Taxonomy. Tephrophilus wetmorei R. T. Moore, 1934, south-east end of Culebrillas Valley, Ecuador. Originally described in a monotypic genus, Tephrophilus; molecular-genetic studies indicate that this species is only distantly related to other members of present genus, and that it should probably either be returned to original genus or placed in enlarged genus containing numerous other mountaintanagers. Monotypic. Distribution. C Andes of SW Colombia (Quindío, and S on both slopes in Cauca and Huila to Nariño), S through Ecuador (mostly on E slope, from Carchi, W Napo and W Morona-Santiago S to E Loja and S Zamora-Chinchipe) and adjacent NW Peru (along Piura–Cajamarca border). Descriptive notes. 20 cm; two birds 62 g and 63 g. Large, stubby-billed tanager with rather odd plumage pattern. Male has crown and nape yellowish-olive, becoming plain olive-green on back and contrasting yellow on rump; side of head down to foreneck black, forming rather broad, “drooping”, roundish mask, forehead and narrow band more or less surrounding mask yellow (much brighter than crown); lesser and median upperwing-coverts dark violet-blue (often hidden), greater coverts black, edged and more broadly tipped dark violet-blue; flight-feathers black, outer feathers narrowly edged dark green, secondaries and tertials more broadly edged and tinged dark green; tail blackish; underparts bright mustard-yellow, usually forming a peak over central foreneck or even to lower throat, sides, flanks and sometimes belly mixed and stained with dusky and olive; iris dark brown; bill blackish, loewr mandible often grey except for blackish tip; legs dark grey. Female is similar to male, but slightly duller. Juvenile apparently undescribed. VOICE. Notably quiet. Infrequently heard song a rather long series of highpitched “tsee” notes, variable in intensity, and remarkably weak for such a large bird; reminiscent of a hummingbird (Trochilidae). Rather low, husky “juur-juur-juur-jur”, possibly as alarm or contact, on taking flight. Habitat. Low mossy forest and very humid elfin woodland, bamboo, and giant grasses at or near tree-line, and in islands of mossy scrub woodland surrounded by grass; at 2900–3650 m. Food and Feeding. Fruit and apparently some insects. Contents of one stomach a large number of seeds. Found singly, in pairs and in little groups up to four individuals, usually seen when accompanying mixed-species flocks containing other tanagers, Buarremon brush-finches etc. Tends to forage mostly within cover, and apt to be seen first as it perches up to fly across small opening. Hops sluggishly as it searches for fruit and other items; once seen to sally clumsily after a flying insect. Breeding. No information. Movements. No information. Status and Conservation. VULNERABLE. Restricted-range species: present in Central Andean Páramo EBA. Generally rare to uncommon; localized. Estimated global population not exceeding 5000 individuals, and probably declining. In Colombia, found at S end of C Andes on both slopes in Cauca and Huila, and recently reported farther N in department of Quindío, also recent sight records in Nariño at Volcán Galeras (SW of Pasto, at 3450 m); present in Puracé National Park and its environs, in Cauca. In Ecuador, found locally near tree-line, mostly on E slope, in Carchi, W Napo, W Morona-Santiago, Azuay, E Loja and S Zamora-Chinchipe; fairly common in Cajanuma sector of Podocarpus National Park, in Loja. In adjacent part of N Peru, recorded from Cerro Chionguela, in E Piura, S along Piura–Cajamarca border. The scatter of records suggests that this species’ distribution is local, although much of the habitat within its range either is not readily accessible to investigators or, where it is accessible, has been subject to extensive deforestation from firewood-cutting, potato cultivation and human settlement. In recent years, it has been found at a number of additional sites in Ecuador but, as yet, at only few sites in Colombia. In both of those countries one of main threats is continued burning of páramo grasslands, which ultimately damages and reduces tree-line and ecotone shrubbery favoured by this tanager; in severely damaged areas tree-line has been burnt back and lowered by several hundred metres in elevation. Habitat loss in its SW Colombian range particularly severe, with estimates of no more than 10% of original tree-line habitat remaining; upper montane forest being replaced by exotic pine (Pinus) plantations near recently discovered site in Nariño. In addition to Podocarpus National Park, which possibly holds a major proportion of the total population, and Puracé National Park, which is regularly burnt, this species has been recorded in at least five other protected areas, most in Ecuador.

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SPECIES ACCOUNTS

PLATE 9

Bibliography. Álvarez-Rebolledo et al. (2007), Anon. (2010a), Arbeláez-Cortés & Baena-Tovar (2006), Butchart & Stattersfield (2004), Clements & Shany (2001), Collar, Crosby & Stattersfield (1994), Collar, Gonzaga et al. (1992), Fjeldså & Krabbe (1990), Hellmayr (1936), Hilty (1985), Hilty & Brown (1986), Isler & Isler (1999), Krabbe et al. (2001), Moore (1934b), Parker et al. (1985), Remsen et al. (2010), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Schulenberg et al. (2007), Sedano & Burns (2010), Stattersfield & Capper (2000), Strewe & Kreft (1999).

Genus ANISOGNATHUS

Reichenbach, 1850

86. Santa Marta Mountain-tanager Anisognathus melanogenys French: Tangara des Santa Marta Spanish: Tangara de Santa Marta German: Schwarzwangen-Bergtangare Other common names: Black-cheeked Mountain-tanager Taxonomy. Poecilothraupis melanogenys Salvin and Godman, 1880, near San Sebastian, 8000 feet [c. 2440 m], Santa Marta Mountains, Colombia. Formerly placed with A. lacrymosus and A. igniventris in a separate genus, Poecilothraupis. May form a superspecies with A. lacrymosus, and in the past treated as conspecific. Monotypic. Distribution. Santa Marta Mts, in N Colombia. Descriptive notes. 18 cm; 41 g (1 male). Strong-billed tanager with mainly blue-andyellow plumage. Has crown and nape shining blue, lores, side of head and neck black (not sharply set off from adjacent grey-blue on side of neck), small yellow spot immediately below eye; upperparts dull dark grey-blue, tail dusky, tinged and edged dark blue; lesser and median upperwing-coverts dull dusky blue, greater coverts dusky, obscurely edged and tipped greenish-blue, flight-feathers and tertials dusky, obscurely edged greenish-blue; throat and entire underparts to undertail-coverts golden-yellow, usually with small amount of black mottling visible on sides and flanks; thigh black; iris dark reddish-brown; bill and legs blackish. Differs from similar A. lacrymosus in having crown blue and underparts bright yellow. Sexes similar. Juvenile apparently undescribed. VOICE. Calls very high-pitched, thin “tic” and “ti” and “ee” notes, sometimes accelerating into trills or rapid bursts of notes. Habitat. Mossy wet-forest borders, second-growth woodland, and regenerating second growth in pastures; less numerous inside wet montane forest. Found over wider range of habitats than is A. lacrymosus. At 1500–3200 m, seasonally as low as 1200 m. Food and Feeding. Known to take a variety of small fruits, and probably also some insects; no other details of diet available. Occurs in pairs and in varyingly sized groups of up to about six individuals, often at fruiting trees and shrubs; often joins mixed-species flocks. Forages from eye level up to canopy, but most foraging in subcanopy or below. Generally conspicuous; runs and hops along mossy branches and into foliage. Breeding. Ten breeding-condition birds in Jan–Jun and Sept; one individual seen while building open cup-nest 14 m up in tree. No other information. Movements. Part of population may shift to lower elevation during peak of rainy season, from Jun through Sept. Status and Conservation. Not globally threatened. Restricted-range species: present in Santa Marta Mountains EBA. Common within its very small global range. Entire population of this species is limited to montane regions in Santa Marta Mts of N Colombia. Deforestation widespread in much of its range, even within Sierra Nevada de Santa Marta National Park, where it occurs. Readily utilizes various types of second growth and edge habitat, however, and is therefore buffered to some extent from forest losses. Numbers may have declined because of deforestation and human settlement, but at present this species unlikely to face serious risk. Bibliography. Donegan & Avendaño (2010), Dunning (1982), Fjeldså & Krabbe (1990), Hellmayr (1936), Hilty & Brown (1986), Isler & Isler (1999), Krabbe (2008), Meyer de Schauensee (1964, 1966, 1970a), Remsen et al. (2010), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Sedano & Burns (2010), Sibley & Monroe (1990), Strewe & Navarro (2004), Todd & Carriker (1922).

87. Lacrimose Mountain-tanager Anisognathus lacrymosus French: Tangara larmoyant

German: Tränenbergtangare

Spanish: Tangara Lacrimosa

Taxonomy. Tachyphonus lacrymosus du Bus de Gisignies, 1846, Maraynioc, Peru. Formerly placed with A. melanogenys and A. igniventris in a separate genus, Poecilothraupis. May form a superspecies with A. melanogenys, and in the past treated as conspecific. Nine subspecies recognized. Subspecies and Distribution. A. l. pallididorsalis Phelps, Sr & Phelps, Jr, 1952 – Perijá Mts on N Colombia–W Venezuela border. A. l. melanops (Berlepsch, 1893) – Andes of W Venezuela (in Trujillo, Mérida and N Táchira). A. l. tamae (Phelps, Sr & Gilliard, 1941) – Andes of W Venezuela (S Táchira) and adjacent E Andes of Colombia (Norte de Santander and Boyacá). A. l. olivaceiceps (Berlepsch, 1912) – Colombia at N end of W Andes (Antioquia) and in C Andes (S to Quindío and Tolima). A. l. yariguierum Donegan & Avendaño, 2010 – Colombia on W slope of E Andes in Santander (Serranía de Yaraguíes). A. l. intensus Meyer de Schauensee, 1951 – SW Colombia on E slope of W Andes (Valle and Cauca). A. l. palpebrosus (Lafresnaye, 1847) – S Colombia (on both slopes in S Cauca and Nariño) and E slope of Ecuador (S to at least Morona-Santiago). A. l. caerulescens (Taczanowski & Berlepsch, 1885) – E slope of Andes in Ecuador (from Loja, or possibly farther N) and N Peru (S to Cajamarca and N Amazonas).

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A. l. lacrymosus (du Bus de Gisignies, 1846) – E slope in Peru from Amazonas (S of R Marañón) S to Junín, Ayacucho and N Cuzco). Descriptive notes. 16–17 cm; 25–38 g. Fairly distinctive tanager with rather short and thick bill. Nominate race has crown and upperparts slaty blue, rump slightly more bluish, tail dusky, obscurely edged blue; side of head down to throat and neck dusky, small but prominent pale yellow spot beneath eye (like teardrop); bend of wing and lesser upperwing-coverts shining cornflower-blue (usually concealed), median and greater coverts dusky, edged blue, flight-feathers dusky, narrowly and obscurely edged greenish-blue, tertials dusky, obscurely edged blue; throat and underparts to undertailcoverts ochre-yellow, thigh olive-yellow, sides of chest and upper breast sometimes with some dusky mottling; iris dark brown; bill blackish; legs slate-grey to black. Sexes similar. Juvenile is duller and paler above and duller yellow below than adult, has olive wash on sides. Races differ mainly in colour or tone of upperparts and underparts, also in head pattern, nominate lacking postauricular spot: intensus is darker above than nominate, deeper ochre below, has both small yellow spot beneath eye and larger yellow spot on rear earcoverts; palpebrosus is very similar to previous, differs in having flight-feathers edged with bluish (not bluish-green), side of head paler, more greenish-slate, and underparts bright warm yellow; caerulescens is intermediate between preceding two, and sometimes lacks yellow spot below eye; olivaceiceps differs from intensus in slightly paler upperparts, and in distinct olive tinge on side of head, side of neck and side of chest; pallididorsalis is distinctive, has forecrown and side of head smoky yellow-olive, crown to back dull bluish-grey (paler than other races), greater wing-coverts slightly paler blue, flight-feathers and tertials narrowly edged paler blue, underparts ochre-yellow, olive mottling on sides and flanks, often lacks yellow spot below eye; tamae is like last, but with darker back, olive mottling continuing down sides and flanks, and normally has yellow spot below eye; recently described yariguierum differs from preceding two in having darker and bluer crown, darker mantle, darker blue rump and shoulder, and darker face with less strong yellowish tones, also darker tail with less extensive blue; melanops is bluer above and golden-yellow to tawnyyellow below, yellower on belly. VOICE. Dawn song in Peru, from exposed perch in top of shrub or small tree, 2–6 phrases composed of high, sharp squeaky notes, e.g. “chuck-zit zit, swick-ee-deeit-it”, or “swickee-deee-deee”, and so on, frequently the same phrase repeated one to several times; resembles song of Thraupis cyanocephala. Possible short song in S Colombia described as an excited “ee-chut-chut-ee”, over and over. High thin “see” or “seeek” notes during foraging; also short bursts of staccato chipping (2–8 notes) over and over. Habitat. Humid and wet montane forest, shrubby forest borders and, at high elevations, stunted mossy forest up to tree-line and locally in patches of woodland and Polylepis above tree-line. At c. 2100–3300 m in most areas, 1800–3500 m in Peru, and as high as 3800 m in Colombia; also recorded down to 1800 m in Venezuela. Food and Feeding. Mostly fruit and insects. Noted as eating berries from melastomes and from Schefflera in Peru. Of 17 stomachs examined, 15 contained only vegetable matter and one only animal matter, and one contained both; contents were fruit pulp, seeds and insects. A conspicuous tanager, often notably confiding; regularly pauses to rest quietly in open or semi-open on low or high branch, sometimes remaining there for several minutes. Pairs or small groups habitually associate with fast-moving mixed flocks containing tyrant-flycatchers (Tyrannidae), emberizid finches and other tanagers; also forage away from them, peering around in leisurely manner. Often forages in large bushes and small trees, and also ascends into subcanopy; ranges over large area. Hops rather heavily in foliage, leaning forward to take fruits, and may glean some insects from leaves, petioles, lichen and bark; occasionally sallies awkwardly for flying insects. Breeding. In Colombia, one nest found in May and begging juvenile in Jul in Cauca (Puracé National Park), and twelve birds in breeding condition in Feb–Aug in Andes. No other information. Movements. Apparently resident. Status and Conservation. Not globally threatened. Fairly common to common; widely distributed in Andes from Venezuela S to S Peru. While deforestation in montane regions has fragmented and reduced the amount of habitat available to it, this species appears not to be faced with any immediate threats. In addition, it is found in many protected areas, including Guaramacal, Sierra Nevada and Tamá National Parks (Venezuela), Los Nevados, Nevado del Huila, Puracé, Paramillo, Las Orquídeas and Farallones de Cali National Parks and Tambito Nature Reserve (Colombia), CayambeCoca, Antisana, Podocarpus and Sumaco-Galeras National Parks (Ecuador), and Tingo Maria National Park and Cordillera de Colán Reserve (Peru). Bibliography. Álvarez-Rebolledo & Córdoba-Córdoba (2002), Donegan & Avendaño (2010), Donegan & Dávalos (1999), Fjeldså & Krabbe (1990), Hellmayr (1936), Hilty (2003), Hilty & Brown (1986), Isler & Isler (1999), Krabbe et al. (2001), Meyer de Schauensee (1964, 1966, 1970a), Meyer de Schauensee & Phelps (1978), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Schulenberg et al. (2007), Sedano & Burns (2010), Sibley & Monroe (1990), Taczanowski (1884), Walker (2001), Weske (1972), Zimmer (1944).

88. Scarlet-bellied Mountain-tanager Anisognathus igniventris French: Tangara à ventre rouge Spanish: Tangara Ventriescarlata German: Mennigohr-Bergtangare Other common names: Black-bellied Mountain-tanager (lunulatus) Taxonomy. A.[glaia] igniventris d’Orbigny and Lafresnaye, 1837, Apolobamba, Bolivia. Formerly placed with A. melanogenys and A. lacrymosus in a separate genus, Poecilothraupis. Four subspecies recognized. Subspecies and Distribution. A. i. lunulatus (du Bus de Gisignies, 1839) – Andes of W Venezuela (S Táchira) and adjacent Colombia (Norte de Santander S in E Andes to Cundinamarca). A. i. erythrotus (Jardine & Selby, 1840) – C Andes of Colombia (Caldas, Cauca, Huila, and both slopes in Nariño) S through Ecuador to N Peru (N of R Marañón, S to R Chamaya). A. i. ignicrissa (Cabanis, 1873) – E slope of Andes of Peru S of R Marañón (C Cajamarca S of R Chamaya, and from Amazonas S to Junín). A. i. igniventris (d’Orbigny & Lafresnaye, 1837) – E slope of Peru (from Junín) S to Bolivia (Santa Cruz). Descriptive notes. 16 cm; 21·5–41 g. Boldly marked tanager with unmistakable colour pattern; bill short and thick. Nominate race has head, throat, chest and upperparts slaty black, mantle and upper

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back slightly glossed blue-black, lower back and rump shining blue (conspicuous in flight); tail slaty black; orange-red triangular patch just behind rear ear-coverts; bend of wing and lesser and median upperwing-coverts shining cornflower-blue, rest of wing blackish; breast, sides and upper belly orange-red, band stretching from upper rear flanks down to thighs and vent black, with variable bluish sheen; iris dark brown; bill and legs black. Sexes similar. Juvenile has black of plumage duller and red replaced by dull brownish-orange to dull orange-rufous; older stages have the orange-rufous stained with red. Races differ mainly in coloration or tone of plumage: erythrotus differs from nominate in having head to back and down to chest deep black (not slaty), triangular neck patch, breast, sides and upper belly bright red (not orange-red), vent area and undertail-coverts solid black; lunulatus differs little from previous, although on average slightly more orange-red, and vent area mixed red and black; ignicrissa is intermediate in colour of red areas, with auricular spot, breast, sides and upper belly brighter than nominate but duller than erythrotus, and lower underparts mixed red and black. VOICE. Sings more than congeners. Song geographically variable; in Venezuela and Colombia frequently at dawn, or at long intervals during the day, a tinkling (like little bells) jumble of rapid, complex, rising and falling notes mixed with lower-pitched nasal notes, somewhat comparable to sound made by cranking of an old engine. Songs of S two races (ignicrissa and nominate) differ from those of N birds but also are variable; typically include numerous squeaky, nasal and mechanical-sounding warbles, with many repeated phrases. Sings mostly from exposed perches at dawn, more from concealment during day. Habitat. Occurs both inside and along borders of montane forest and elfin woodland, also in thick hedgerows, shrubby hillsides, second growth and disturbed or agricultural areas and remnant patches of woodland up to edge of páramo and puna; occasionally in parks and gardens and green belts in settled areas. All races utilize wide range of habitats, but S ones seem more closely associated with forest, those in N of range more often at forest borders or in semi-open areas, woodlots, and lines of trees and shrubs along fencerows. Mostly at 2600–3500 m; rarely down to 2250 m and up to as high as 3900 m. Food and Feeding. Eats mostly fruit, also petals or centres from flowers, and nectar; also insects. Of 44 stomachs examined, 41 contained only vegetable matter, one held only animal matter and two contained both; contents were fruit pulp, seeds, leaves, buds, and insects. Occurs in pairs or in loosely associated groups of up to about eight, occasionally more, which forage alone or join mixed flocks of other tanagers and finches in forest or along forest borders. Often seen as it moves in somewhat undulating and usually low flight across openings between hedgerows or forest patches, but once in forest can be difficult to see. Forages from low to high levels, e.g. average foraging height in Bolivia was c. 6 m (mostly in upper third of trees); may sit relatively immobile near top of or in inner part of a tree for a moment, and then hop through foliage. Fruit generally taken by reaching outwards or downwards, without acrobatics. Gulps or mashes fruit; may mash flowers for nectar. In Bolivia, observed obtaining insects by reaching under, hanging down, or sallying 2 m out to air. Breeding. Fledglings in Feb, Apr and Jun in Colombia, Jan and Jun–Sept in Ecuador, Dec in Peru and Mar in Bolivia; in Colombia, one bird in breeding condition in Nov, three in late Dec, six in Feb and two in Jul. Strongly territorial in Peru. Nest found in trees and thickets in Colombia. Eggs presumed of this species were pale greenish-white with small reddish-brown and violet-grey spots, especially at large end. No other information. Movements. No local or seasonal movements verified, but wanders widely and regularly appears in parks and other green areas in cities (e.g. on Sabana de Bogotá, in Colombia) if these close to forest. Individuals seen well above or below normal altitudinal range may be wanderers or represent elevational movements. Status and Conservation. Not globally threatened. Uncommon to locally common. Despite extensive deforestation within parts of its range, this species is able to utilize a broad spectrum of natural and disturbed or man-altered habitats, preventing it from suffering severe impacts from loss of primary forest. Occurs also in numerous parks and other protected areas, as well as unprotected ones, and thrives locally in many heavily intervened areas so long as some woodlots, patches of forest and shrubby fencerows remain. Occurs in Tamá National Park (Venezuela), Sierra Nevada del Cocuy, Chingaza and Puracé National Parks and many others (Colombia), Cayambe-Coca, Sangay and Podocarpus National Parks (Ecuador), Cordillera Azul, Tingo Maria and Manu National Parks (Peru), and Isiboro Sécure and Amboró National Parks (Bolivia). Bibliography. Álvarez-Rebolledo & Córdoba-Córdoba (2002), Anon. (2000), Barat (1980), David & Gosselin (2002a), Dunning (1982), Fjeldså & Krabbe (1990), Hellmayr (1936), Hilty (2003), Hilty & Brown (1986), Isler & Isler (1999), Krabbe et al. (2001), Meyer de Schauensee (1966, 1970a), Meyer de Schauensee & Phelps (1978), Moynihan (1979), Ogilvie-Grant (1912), Olivares (1969), Peters & Griswold (1943), Remsen et al. (1987), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Schulenberg et al. (2007), Sedano & Burns (2010), Walker (2001), Zimmer (1944).

89. Blue-winged Mountain-tanager Anisognathus somptuosus French: Tangara somptueux German: Blauflügel-Bergtangare Spanish: Tangara Aliazul Other common names: Blue-shouldered/Blue-rumped Mountain-tanager Taxonomy. Tachyphonus somptuosus Lesson, 1831, no locality = Peru. This species, A. flavinucha and A. notabilis were formerly placed in a separate genus, Compsocoma. Usually treated as conspecific with A. flavinucha, but the two differ markedly in voice and to lesser degree also in plumage; their ranges are believed to be geographically isolated, although approaching each other closely. Present treatment as separate species provisional, pending genetic and further vocal analysis of the various taxa. Race alamoris extremely similar to nominate and possibly better synonymized with it; provisionally retained pending thorough review of all races. Eight subspecies currently recognized. Subspecies and Distribution. A. s. venezuelanus (Hellmayr, 1913) – N Venezuela (coastal cordillera from Yaracuy E to Miranda). A. s. virididorsalis (Phelps, Sr & Phelps, Jr, 1949) – Aragua (Cerro Golfo Triste), in N Venezuela. A. s. antioquiae (Berlepsch, 1912) – N end of W Andes, both slopes of C Andes (in Antioquia) and E slope (in Tolima), in N Colombia. A. s. victorini (Lafresnaye, 1842) – Andes of W Venezuela (S Táchira) and W slope of E Andes of Colombia (from Santander S to head of Magdalena Valley, in Huila).

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A. s. cyanopterus (Cabanis, 1866) – Andes of S Colombia (W slope of C range S from Quindío) and W Ecuador (S to Chimborazo and NE Azuay). A. s. alamoris (Chapman, 1925) – S Ecuador (W slope in S Azuay, El Oro and W Loja). A. s. baezae (Chapman, 1925) – E slope of Andes in SW Colombia (Nariño) and Ecuador (S to Morona-Santiago). A. s. somptuosus (Lesson, 1831) – E slope of Andes in extreme S Ecuador (Zamora–Chinchipe) S to C Bolivia (La Paz, Cochabamba). Descriptive notes. 16–17 cm; 33–56 g (combined weight data from when lumped with A. flavinucha). Distinctive strong-billed tanager with mostly blue wings and bold plumage pattern. Nominate race has forehead and side of head to well below eye black, fairly broad yellow stripe from centre of crown to nape (stripe sometimes partly concealed); mantle and back deep black, rump dark brownish-olive becoming greener on uppertailcoverts; tail blackish, at least several outer pairs of feathers edged bright blue; marginal coverts and lesser and median upperwing-coverts shining cobalt-blue, greater coverts black; primary coverts black, flight-feathers black, broadly edged bright turquoise-blue, tertials deep black; throat and entire underparts bright yellow except for black rear thigh, underwing-coverts bright yellow; iris dark reddish-brown; upper mandible blackish, lower mandible bluish-grey with blackish tip and base; legs dark horn-grey. Differs from similar A. notabilis mainly in much darker, black mantle and back. Sexes similar. Juvenile has yellow in plumage duller. Races vary mainly in colour and pattern of upperparts: baezae differs from nominate in having back and rump mixed black and olive; alamoris is very like nominate (with same turquoise-blue flight-feather edging), but perhaps slightly smaller and with upperparts slightly tinged green; cyanopterus is much like nominate, but rump dark olive-green (not dark brownish-olive) and wing edgings darker cobalt-blue (not bright turquoise-blue); victorini is slightly larger, and has mantle to uppertail-coverts moss-green; antioquiae has centre of mantle and back tinged greenish, rump green, wing and tail edged medium blue; virididorsalis differs in having back greenish-black, flight-feathers edged cobalt-blue like shoulder (not bright turquoise-blue), and tail feathers edged cobalt-blue; venezuelanus differs from nominate in slightly smaller size, back sooty black (duller than nominate), and lower rump and uppertail-coverts sooty black with yellowish-green tinge. VOICE. Generally rather quiet. Song a weak, rapid, very high-pitched, almost twittering “ti-ti-ti-ti ti’ti’ti’tí’tí’títi’ti’ti’ti’ti’ti’ti ti ti”, a little louder and faster in middle, in N & C Peru described as a high, weak wheedling “si-titi-si-titi tsu-ti-su-ti-tsu-ti-tsu-ti”; very unlike song of A. flavinucha. When foraging, gives soft “tic” and “teep” notes, or short bursts of “tic” notes, sometimes extended into little trills. Habitat. Humid and wet forest, forest borders, and tall second-growth woodland. At 900–2100 m in Venezuela, mostly 1400–2600 m in Colombia, and 1200–2500 m in Ecuador. Food and Feeding. Insects, and small fruits and berries. Conspicuous pairs or groups of 3–10 individuals forage alone or join mixed-species flocks typically containing other tanagers as well as New World warblers (Parulidae), but frequently strike out on their own in unpredictable or erratic little forays. Forages at various heights, from below eye level to (occasionally) high in canopy, and inside or at edge of forest; median foraging height in Valle (Colombia) was c. 12 m. Energetic, foraging actively in distinctive, stereotyped manner, nimbly run-hopping along slender, mostly bare branches, working out fairly quickly to terminal foliage clusters, where it pauses or hops deliberately and spends up to 30 seconds in peering around carefully, stretching up, leaning down, and occasionally pecking or lunging short distance for prey, sometimes sallying or hovering; then flies to another branch and repeats process. Also regularly checks hanging dead leaves for insects. Takes Miconia berries mostly without acrobatics. Breeding. In Colombia, nests in Apr–Jun in SW (La Planada Nature Reserve), nestlings in Jun and fledglings in Nov in Valle del Cauca, and juveniles in Mar and Sept–Oct, and three breedingcondition birds in Mar and 17 more May–Sept in N; in Ecuador, juveniles in Jan in S and adult carrying nest material in early Mar in SW (Tambo Negro, in Loja, 600–1000 m); juveniles in Mar in Peru. Co-operative breeding, with four adult helpers, recorded in Colombia. At La Planada (Colombia), nest an open cup 12·4 cm in diameter and 8·6 cm high, made from soft plant material including rootlets, green moss, fern leaves and fine vegetable fibres, lined with finer moss and dry bamboo leaves, one placed 3·5 m up on horizontal limb of pepper (Piper) covered with tree-fern leaves and bamboo at disturbed forest border, another 6·5 m up on horizontal branch of Clusia, and a third 7·5 m up; in captivity, cup-nest of coarse grass and lined with finer grass. Following details taken from S Colombia (La Planada): at one nest one dominant individual (possibly female) and four adult helpers were present, helpers fed dominant bird, which in turn fed the two nestlings, as also did helpers, two fledglings subsequently seen with the five adults c. 50 m from nest tree and all seven remained together for c. 7 months or possibly more; at a second nest, one bird incubated and four helpers present; a third nest was attended by only two adults, which were seen to carry food to nest. Movements. Apparently resident. Status and Conservation. Not globally threatened. Fairly common, and widespread across most of its large Andean range. Locally may occur at low density. Present in many protected areas, e.g. Henri Pittier, Guaramacal, Sierra Nevada and Tamá National Parks (Venezuela), Chingaza, Cueva de los Guácharos, Nevado del Huila, Puracé, Las Orquídeas, Farallones de Cali and Munchique National Parks anmd La Planada and Tambito Nature Reserves (Colombia), Sangay, CayambeCoca and Podocarpus National Parks (Ecuador), and Cordillera Azul and Tingo Maria National Parks (Peru). Occurs also in many unprotected areas with suitable habitat. Andean deforestation has resulted in local range contraction and fragmentation. Bibliography. Álvarez-Rebolledo & Córdoba-Córdoba (2002), Best et al. (1996), Burns (1997a), Burns & Racicot (2009), Donegan & Dávalos (1999), Fjeldså & Krabbe (1990), Hellmayr (1936), Hilty (2003), Hilty & Brown (1986), Isler & Isler (1999), Krabbe & Nilsson (2003), Mee et al. (2002), Meyer de Schauensee (1966), Meyer de Schauensee & Phelps (1978), Miller (1963), Moore et al. (1999), Norgaard-Olesen (1973), Phelps & Phelps (1950), Remsen et al. (2010), Restall et al. (2006), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman, Donegan & Caro (2008), Salaman, Donegan & Cuervo (1999), Schäfer & Phelps (1954), Schulenberg (2000b), Schulenberg & Plenge (1980), Schulenberg et al. (2007), Sedano & Burns (2010), Sibley & Monroe (1990), Strewe (2001a, 2001b), Walker (2001), Zimmer (1944).

90. Bolivian Mountain-tanager Anisognathus flavinucha French: Tangara à nuque jaune Spanish: Tangara Boliviana German: Blauschwingen-Bergtangare

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SPECIES ACCOUNTS Taxonomy. T.[achyphonus] flavinucha d’Orbigny and Lafresnaye, 1837, Yungas, Bolivia. This species, A. somptuosus and A. notabilis were formerly placed in a separate genus, Compsocoma. Usually treated as conspecific with A. somptuosus, but the two differ markedly in voice and to lesser degree in plumage; their ranges are believed to be geographically isolated, although approaching each other closely. Present treatment as separate species provisional, pending genetic and further vocal analysis of the various taxa. Monotypic. Distribution. E slope of Andes in Peru (from Puno) S to S Bolivia (Chuquisaca). Descriptive notes. 16–17 cm; 33–56 g (combined weight data from when lumped with A. somptuosus). Has head, throat, mantle and back deep black, prominent bright yellow median crownstripe on hindcrown and nape; lower back and rump bright cobalt-blue; lesser and median upperwing-coverts purplish-blue, greater coverts black, primary coverts, flightfeathers and tail black, feathers sharply edged pale turquoise-blue, tertials mostly black; underparts bright golden yellow; iris dark brown; bill black, base of lower mandible blue-grey; legs dark grey. Differs from very similar A. somptuosus mainly in having more restricted yellow crownstripe, and cobalt-blue (not olive) rump. Sexes similar. Juvenile is much like adult, but duller yellow below, with yellow feathers of rear crown and nape mixed or tipped with black. VOICE. Frequently heard song, at dawn and in morning hours, while perched or in flight display as bird climbs 30–45 m on shallowly beating wings, a series of musical whistles, soft but gradually increasing in volume and pitch, e.g. “too-too tyoo-towoo-towoo too-wít too-wít too-wít”, rather loud at end; in S Peru described as a rising series of wiry, quavering notes, “tchu-tchu-twéé-aww twéé-awéé wéé-wéé wéé-wéé”; dramatically different from song of A. somptuosus. Calls are high “tic” notes. Habitat. Humid and wet montane forest, forest borders and older second growth, generally not away from taller forest. At c. 1500–2500 m. Food and Feeding. Insects and fruit; takes large melastome fruits. Forages mainly in middle levels up to canopy; sometimes descends into shrubbery in undergrowth of forest or along forest edges. Hops along mossy epiphyte-covered branches away from leaves, and searches moss and bark. Behaviour much like that of A. somptuosus. Breeding. In Bolivia, bird in breeding condition reported in Aug and eggs in Oct in La Paz, and fledglings in Apr in Cochabamba. No other information. Movements. Apparently resident. Status and Conservation. Not globally threatened. Uncommon to locally fairly common. Range much smaller than that of allied A. somptuosus, but montane habitat in S Peru and adjacent Bolivia remains in relatively good shape. Occurs in several large protected areas, including Manu National Park, in Peru, and Madidi, Isiboro Sécure and Amboró National Parks, in Bolivia. Bibliography. Bleiweiss (2004), Clements & Shany (2001), Fjeldså & Krabbe (1990), Hesse (1980a), Isler & Isler (1999), Krabbe & Nilsson (2003), Mayer (2000), Norgaard-Olesen (1973), Ridgely & Tudor (1989, 2009), Schulenberg et al. (2007), Weske (1972), Zimmer (1944).

91. Black-chinned Mountain-tanager Anisognathus notabilis French: Tangara à menton noir Spanish: Tangara Barbinegra German: Schwarzkinn-Bergtangare Taxonomy. Tanagra notabilis P. L. Sclater, 1855, eastern range of cordilleras north of Quito, Ecuador. This species, A. somptuosus and A. flavinucha were formerly placed in a separate genus, Compsocoma. Monotypic.

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PLATE 9 Distribution. W Andes of Colombia (Pacific slope from headwaters of R San Juan, in Risaralda) S to W Ecuador (El Oro and W Loja). Descriptive notes. 18 cm; one male 44·1 g, one female 68·5 g. Quiet tanager with svelte plumage and fairly strong and rather pointed bill. Has forecrown, side of head, nape and chin black, broad median stripe on rear crown bright yellow; upperparts bright yellow-olive, becoming olive-yellow on rump; tail entirely black (or feathers very obscurely edged dark violetblue); lesser and median upperwing-coverts plain olive, greater coverts blackish, broadly edged violet-blue, flight-feathers black, narrowly edged dark violet-blue, tertials black, narrowly edged violet-blue; central throat and underparts, including undertail-coverts, rich orange-yellow, underwing-coverts bright yellow; iris dark red; bill black, bluish-grey on base of lower mandible; legs black. Distinguished from A. somptuosus mainly by olive (not black) back, less conspicuous wing edgings and richer orange (not yellow) underparts. Sexes similar. Juvenile apparently undescribed. VOICE. Not particularly vocal, even when foraging. Infrequently heard song a series of high-pitched notes, “tsit-tsit-tseeéu-tsit-tseeu-tsit”, repeated over and over; calls high thin “tic” notes. Habitat. Humid and wet mossy forest and forest borders. At 900–2200 m (once to 300 m) in Colombia; mostly 1400–2200 m in Ecuador, but in El Oro as low as 800–1100 m. Food and Feeding. Fruit; presumably also insects. Nestlings in Colombia were fed with relatively large flying insects, including beetles (Coleoptera), hymenopterans and lepidopterans, and fruits (Alchornea, Clusia and Miconia identified). Usually seen singly or in pairs, much less often in groups up to six individuals; singles, pairs or groups often forage with mixed-species flocks. May sit more or less still and rather erect, then briskly run-hop along branches and peer at foliage or fruit at various heights, from fairly low along wooded borders to much higher in taller forest. During brood-feeding period, captures many insects in air in short hawking manoeuvres, 5–20 m from branches. Generally less conspicuous than A. somptuosus, although the two are sometimes found together. Breeding. In SW Colombia, three nests found during 10th Apr and 5th May at La Planada Nature Reserve (in Nariño); following details refer to these. Nest constructed within clump of moss and epiphytes 13·4–16·5 m above ground on tree limb in undisturbed wet, epiphyte-laden montane forest on ridge (at 1740–1780 m). First nest attended by three adults, all carrying food to nestlings (insects captured generally within c. 50 m of nest tree); after this nest failed, one observed carrying moss and dry leaves to new site, and by end May all three were again feeding young; seven days later, three fledglings seen near nest-site; inferred incubation period c. 12–14 days and nestling period c. 13–15 days. No breeding-territory overlap with A. somptuosus, which favoured forestedge sites. Movements. Apparently resident. Status and Conservation. Not globally threatened. Restricted-range species: present in Chocó EBA. Uncommon to locally fairly common in Ecuador; generally less numerous and more local in Colombia. In general less numerous and less conspicuous than A. somptuosus. Deforestation, various farming activities and illegal cultivation of coca and marijuana within this species’ range (especially in SW Colombia) are ongoing threats within the rather narrow elevational band of Pacific-slope montane forest where it occurs. Present in several protected areas, including Farallones de Cali and Munchique National Parks and La Planada, Tambito and El Pangan Nature Reserves (Colombia), and El Angel and Cotocachi-Cayapas Ecological Reserves and Maquipucuna Reserve (W Ecuador). Although populations trends for this species have not been quantified, it is almost certainly declining. Bibliography. Bleiweiss (2004), Donegan & Dávalos (1999), Hilty & Brown (1986), Isler & Isler (1999), Meyer de Schauensee (1951, 1952b, 1964, 1966, 1970a), Moore et al. (1999), Orejuela & Cantillo (1990), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Sedano & Burns (2010), Strewe (2001a, 2001b).

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ssp riefferii ssp carrikeri

ssp taeniata

ssp stictocephala

ssp bolivianus

95 98

96 97

99 100 ssp caeruleoventris ssp rufivertex

variants

101 ssp venezuelensis

ssp melanonota

102 ssp bourcieri

103 ssp calliparaea

104

PLATE 10 inches

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ssp fulgentissima

206

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Family THRAUPIDAE (TANAGERS)

207

SPECIES ACCOUNTS

Genus CHLORORNIS

PLATE 10 Reichenbach, 1850

92. Grass-green Tanager Chlorornis riefferii French: Tangara de Rieffer

German: Papageitangare

Spanish: Tangara Lorito

Taxonomy. Tanagra Riefferii Boissonneau, 1840, Santa Fé de Bogotá, Colombia. Molecular-genetic studies place present species within Buthraupis, most closely related to B. eximia and B. aureodorsalis. Five subspecies recognized. Subspecies and Distribution. C. r. riefferii (Boissonneau, 1840) – Andes of Colombia and Ecuador. C. r. dilutus J. T. Zimmer, 1947 – Andes of N Peru S of Chachapoyas. C. r. elegans (Tschudi, 1844) – Andes of C Peru (Junín). C. r. celatus J. T. Zimmer, 1947 – Andes of S Peru below Limbani. C. r. bolivianus (Berlepsch, 1912) – Andes of Bolivia (La Paz). Descriptive notes. 20 cm; 42–59 g. Large, unmistakable tanager with grass-green plumage, robust proportions and orange bare parts; bill thick and somewhat conical. Nominate race is bright shining green, with narrow area of chestnut on base of forehead, lores, ocular area, cheek and chin (forming conspicuous mask); bend of wing and marginal coverts pale blue (almost always hidden), otherwise upperwingcoverts green, primary coverts dusky, edged green, flight-feathers dusky, heavily edged green, tertials dusky, broadly edged green; undertailcoverts chestnut; iris chestnut-brown; bill bright orange; legs bright orange, claws dusky orange. Sexes similar. Juvenile is much like adult, but duller green and with bill dusky. Races differ relatively little: bolivianus is lighter and brighter green than nominate, has larger chestnut mask extending across entire forehead, side of head and most of throat; dilutus has blue forehead (inconspicuous) and paler chestnut on face; celatus has chestnut of forehead narrowly edged pale blue above; elegans is like last, but with chestnut of chin extending to upper throat. VOICE. Dawn song 1–2 “enk” notes followed by bursts of 1–5 harsh, squeaky, doubled notes like sound of squeaky gate being rapidly moved. Possible day song a heavy “chink-zheet”, second note higher, repeated once every 2 seconds. Most frequently heard vocalization is contact call during foraging, a rather low-pitched, nasal, strained “eenk” or “eck”, not loud, notes given singly, paired or sometimes several times in irregular series; infrequently in long stuttering and louder series, especially just prior to or during flight, e.g. “t-e-e-e-e-e-k”. Habitat. Wet, mossy and often rather low-stature or stunted upper montane forest and forest borders almost to tree-line; occasionally dense, mossy second growth. Recorded at c. 1500–3500 m; 1700–3300 m (mostly 2400–2700 m) in Colombia, mostly 2000–2900 m in Ecuador, and at 2000–3500 m in Peru. Food and Feeding. Of seven stomachs examined, three contained only vegetable matter and two only animal matter, and two held both; contents included fruit pulp, berries, seeds, a 40-mm worm, insect remains and caterpillars. Usually seen in pairs or in small groups of 3–6 individuals, less frequently up to ten; in Bolivia (where apparently less numerous), more often seen singly or in pairs. Regularly forages with mixed-species flocks, then moving sluggishly from branch to branch. Inspects moss and epiphytes with peering movements and, especially, by bending or leaning down to look at mossy sides or undersurfaces of branches 3–6 cm in diameter; may hang head downwards or flutter heavily in foliage. Breeding. Nests found in Jul and Sept in Ecuador. One nest in Colombia was relatively large, constructed from green moss and decorated with ferns; contained 1 egg, grey, freckled lilac-grey. In Ecuador, two nests in consecutive years in Pichincha (near Tandayapa, at 2250 m), second one only 2 m away from first, believed built by same pair, made from moss, ferns and fine rootlets, lined with tree-fern scales, placed beneath moss on steep roadside bank; 1 egg. No other information. Movements. Apparently resident. Status and Conservation. Not globally threatened. Fairly common throughout most of range; rather uncommon in Bolivia. Present in a number of protected areas, including Las Orquídeas, Farallones de Cali, Munchique, Puracé, Cueva de los Guácharos and Chingaza National Parks and Tambito Nature Reserve (Colombia), Sangay and Podocarpus National Parks (Ecuador), Manu National Park (Peru) and Madidi National Park (Bolivia). Considerable intact but unprotected forest remains within this species’ high-elevation range, but in many areas it is subject to firewood-cutting, and to deforestation for expanding pastureland for cattle, various agricultural crops, and human settlement. Bibliography. Burns (1997a), Burns & Naoki (2004), Burns et al. (2002, 2003), David & Gosselin (2002b), Donegan & Dávalos (1999), Dunning (1982), Fjeldså & Krabbe (1990), Goodfellow (1901), Greeney & Gelis (2006b), Hilty & Brown (1986), Isler & Isler (1999), Krabbe et al. (2001), Mitsch (1978), Moore et al. (1999), Niethammer (1956), Remsen (1985), Remsen et al. (2010), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman, Donegan & Caro (2008), Salaman, Donegan & Cuervo (1999), Schulenberg et al. (2007), Sclater & Salvin (1879), Sedano & Burns (2010), Taczanowski (1884), Vuilleumier (1970), Walker (2001), Weske (1972), Zimmer (1947b).

Taxonomy. Tanagra (Tachyphonus) taeniata Boissonneau, 1840, Santa Fé de Bogotá, Colombia. Molecular-genetic data indicate that this genus and Delothraupis are sister-taxa, with Calochaetes sister to both. Races have in the past been treated as three separate species. It was recently proposed that S race stictocephala be regarded as a separate species (carrikeri retained with nominate race), on grounds of known vocal and morphological evidence; this merits further examination. Three subspecies currently recognized. Subspecies and Distribution. D. t. carrikeri Wetmore, 1946 – Santa Marta Mts, in N Colombia. D. t. taeniata (Boissonneau, 1840) – Andes of Venezuela (Trujillo S to Táchira), all three ranges in Colombia (in W Andes mainly N & S ends, in Antioquia and Cauca), and both slopes in Ecuador and NW Peru (S to Piura and N Cajamarca N of Marañón Valley). D. t. stictocephala Berlepsch & Stolzmann, 1894 – E slope of Andes in Peru (from S side of Marañón Valley, in Amazonas, S to N end of Cordillera Vilcanota, in Cuzco). Descriptive notes. 20 cm; 31–45 g. Colourful, sturdy-billed, somewhat retiring tanager often noted first through its distinctive voice. Nominate race has head to nape, upper mantle and foreneck black, feathers of forecrown and superciliary area tipped silvery blue, forming long, broad and rather loosely defined frosty supercilium extending almost to nape and often partly down side of neck; most of mantle to rump dark dusky blue, tail blackish, tinged dark blue; lesser upperwing-coverts broadly edged silvery blue, median and greater coverts dusky, broadly tipped slightly darker blue, greater coverts also broadly edged pale greyish violet-blue; flight-feathers and tertials dusky, broadly edged dull blue, flight-feathers narrowly edged dull blue, tertials more broadly edged dull blue; broad pectoral band buff, varying in intensity from whitish-buff to greyish-buff, lower breast and belly bright yellow, thigh dusky blue, undertailcoverts buff; iris dark reddish-brown; bill black; legs blackish. Sexes similar. Juvenile has dull olive-green back and chest, vent area duller and more ochraceous rather than buff. Race carrikeri is much like nominate, but forecrown and eyebrow more densely streaked, chin mixed and streaked black and buff, darker buff colour on chest and extending up to centre of throat, and breastband slightly darker, more cinnamon-buff; stictocephala has entire crown and nape somewhat streaked and frosted with silvery blue-white, mantle dark blue (like back), buff breastband narrower, and vent and undertail-coverts yellow (not buff). VOICE. Song of nominate race and carrikeri, repeated over and over at short intervals early in morning, a loud, whistled “phéééé-baay” delivered slowly, second note lower, or downslurred, occasionally expanded to 3 notes, e.g. “téééé teeee tyyyy”. In S Peru (stictocephala) a thin, high-pitched, single-note whistle sharply slurred downwards, reminiscent of song of a Pipreola fruiteater; this single-note song also may be continually repeated at dawn, and sometimes later in morning. May sing from exposed medium-height perch, but more often when partly concealed in vegetation, or at edge of concealing vegetation. Habitat. Thick, mossy, wet montane forest and broken forest with dense lower vegetation, especially with Chusquea bamboo thickets, and dense tangles inside forest and in gaps and edges; locally up to elfin woodland at tree-line and in patches of Polylepis and shrubs slightly above treeline. At 2000–3000 m in Venezuela, 2400–3600 m in Colombia, 2500–3500 m in Ecuador, and 1900–3500 m in Peru; in all areas except Venezuela, most numerous above c. 2500 m. Food and Feeding. Small fruits and berries, including mistletoe (Loranthaceae), and insects. Three stomachs contained both vegetable and animal matter, including fruit, seeds and insects. Single individuals or pairs, usually heard before they are seen, follow mixed-species flocks or wander alone over large areas. Rather slow-moving; forages from eye level to treetops, but more often fairly low, and often in dense vegetation, where it keeps out of sight. Usually remains at least partially concealed in foliage, but occasionally sits up in the open on top of bush for a few moments. Searches mossy branches for insects. Hops along large limbs, scans upper surfaces, and bends down to look under limbs or in moss, bark and especially lichens; also observed when searching moss on steep banks. Breeding. Juveniles and immatures in Jan and Sept in NW Ecuador; in Colombia, two breedingcondition birds in late Dec in Boyacá, and a further eight in Feb–Sept in Santa Marta Mts and E & C Andes. No other information. Movements. Apparently resident. Status and Conservation. Not globally threatened. Fairly common almost throughout its range, and widespread. Appears to require fairly large tracts of humid montane forest; has declined or disappeared locally in the face of deforestation, firewood-cutting and human settlement. Can be found in many national parks and other protected areas throughout its range. In addition, extensive unprotected but intact habitat exists within its range, and most of this appears secure at present. Bibliography. Álvarez-Rebolledo & Córdoba-Córdoba (2002), Burns (1997a), Burns & Naoki (2004), Burns et al. (2003), Donegan & Dávalos (1999), Dunning (1982), Fjeldså & Krabbe (1990), Hilty (2003), Hilty & Brown (1986), Krabbe, Flórez et al. (2006), Krabbe, Moore et al. (2001), Meyer de Schauensee (1964, 1966, 1970a), Meyer de Schauensee & Phelps (1978), Olivares (1963), Parker & O’Neill (1980), Remsen et al. (2010), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Schulenberg et al. (2007), Sedano & Burns (2010), Walker (2001), Weske (1972), Zimmer (1944).

Genus DELOTHRAUPIS Genus DUBUSIA

P. L. Sclater, 1886

Bonaparte, 1850

94. Chestnut-bellied Mountain-tanager 93. Buff-breasted Mountain-tanager

Delothraupis castaneoventris

Dubusia taeniata

French: Tangara à ventre marron Spanish: Tangara Ventricastaña German: Braunbauch-Bergtangare

French: Tangara à poitrine fauve German: Silberbrauen-Bergtangare Spanish: Tangara Pechifulva Other common names: Carriker’s Mountain-tanager (carrikeri)

Taxonomy. Calliste castaneoventris P. L. Sclater, 1851, Bolivia.

On following pages: 95. Diademed Tanager (Stephanophorus diadematus); 96. Purplish-mantled Tanager (Iridosornis porphyrocephalus); 97. Yellow-throated Tanager (Iridosornis analis); 98. Golden-collared Tanager (Iridosornis jelskii); 99. Golden-crowned Tanager (Iridosornis rufivertex); 100. Yellow-scarfed Tanager (Iridosornis reinhardti); 101. Fawnbreasted Tanager (Pipraeidea melanonota); 102. Glistening-green Tanager (Chlorochrysa phoenicotis); 103. Orange-eared Tanager (Chlorochrysa calliparaea); 104. Multicolored Tanager (Chlorochrysa nitidissima).

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Family THRAUPIDAE (TANAGERS)

PLATE 10

SPECIES ACCOUNTS

Molecular-genetic data indicate that this genus and Dubusia are sister-taxa. Species is sometimes placed in latter genus, in part because their songs are somewhat similar to each other. Birds from Auquimarca (Junín), in Peru, described as a geographical race, peruviana, but appear to differ little, if at all, from those in rest of species’ range. Treated as monotypic. Distribution. E slope of Andes in Peru (from SE La Libertad) S to Bolivia (La Paz and Cochabamba). Descriptive notes. 15 cm; 25–33 g. A small tanager, mainly dull blue above and rufous below, with small bill. Has crown medium skyblue with slight frosty sheen (pale feather edges), bordered below by narrow black mask from base of forehead and lores back through eye, mask separated from narrow black submalar line by pale malar line mixed rufous and white; upperparts, including uppertail-coverts, rich dark blue, upperwing-coverts dusky, broadly edged and tipped medium blue, flight-feathers, tertials and tail black, edged medium blue; chin whitish, throat and underparts, including undertail-coverts, rufous; iris red to dark red; bill black, basal portion of lower mandible blue-grey; legs blackish. Sexes similar. Juvenile apparently undescribed. VOICE. In general not a persistent singer. Song in Peru a clear, whistled “peee paay”; in Bolivia (possibly also Peru) extended to 3 notes, “peeee pee pay” or “pee pee pay”, first note highest, and song may be repeated at short intervals early in morning. Both 2-note and 3-note songs recall those of Dubusia taeniata, but not so loud or forceful. Also a sharp, high-pitched and slightly descending “tzeeeeee”, possibly also song. Call a brief “zee zee”, like that of many other tanagers. Habitat. Dense humid and wet montane forest, elfin woodland, thickets, and gaps and borders in broken montane forest, and patches of scrubby vegetation at tree-line. Recorded mainly at 2000– 3500 m in Peru, seasonally down to at least 1650 m in Cuzco (S Peru) and in Bolivia. Food and Feeding. Arthropods and berries. Normally found singly or in pairs, infrequently three or occasionally more individuals, alone or, more often, with large mixed-species flocks containing tyrant-flycatchers (Tyrannidae), New World warblers (Parulidae) and other tanagers. Stays mostly in canopy or upper levels, occasionally descending to eye level, even near ground along dense forest borders; median foraging height in Bolivia was c. 10 m, or 2·5 m below top of canopy. Takes berries from a perched position. Searches for insects while hopping in rather deliberate manner along large, moss-covered branches. Takes most insect prey from moss; leans head downwards and occasionally hangs upside-down to probe mossy undersides of branches; also picks prey from both surfaces of leaves and from petioles, and has been noted as sallying to air for small flying insects. Breeding. No information. Movements. Local or seasonal movements believed likely, although none reported. Status and Conservation. Not globally threatened. Uncommon. Distributed along montane and tree-line Andean habitats from N Peru S to C Bolivia. Occurs in various protected areas, including Tingo Maria and Manu National Parks (Peru) and Madidi and Amboró National Parks (Bolivia). Main threats to its habitat are deforestation and firewood-cutting, similar to threats faced by many other montane species. Its range also includes considerable intact, unprotected habitat which appears to be at relatively little short-term risk. Bibliography. Burns (1997a), Burns & Naoki (2004), Burns et al. (2002, 2003), Fjeldså & Krabbe (1990), George (1964), Hilty (2009b), Isler & Isler (1999), Klicka et al. (2007), Meyer de Schauensee (1966, 1970a), Parker & O’Neill (1980), de la Peña & Rumboll (1998), Remsen (1984), Ridgely & Tudor (1989, 2009), Schulenberg et al. (2007), Sedano & Burns (2010), Walker (2001), Zimmer (1944).

Genus STEPHANOPHORUS

Strickland, 1841

95. Diademed Tanager Stephanophorus diadematus French: Tangara à diadème German: Diademtangare Other common names: White-capped Tanager!

Spanish: Tangara Diademada

Taxonomy. Tanagra diademata Temminck, 1823, Curitiba, Paraná, Brazil. Monotypic. Distribution. SE Brazil (Paraná and São Paulo E to S Minas Gerais and Rio de Janeiro, S to Rio Grande do Sul), adjacent E Paraguay, Uruguay and NE Argentina (E from extreme E Santa Fe, S to N Buenos Aires). Descriptive notes. 19 cm; two males 35 g and 41·5 g. Rather robust tanager, colourful in good light but most often appearing blackish, with pale crown patch, feathers of forecrown short, dense and plushy; bill short and thick. Has forecrown, lores, ocular area and chin black, forming small mask; large white crown patch surmounted by small but fairly conspicuous red tuft at front (both white and red feathers can be erected); rest of plumage mostly shining dark blue, upperwing-coverts, flight-feathers and tail feathers blackish with heavy dark blue edges (blue edging inconspicuous); iris dark brown; bill blackish; legs dusky grey. Sexes similar. Immature is much duller and duskier than adult, with features of crown obscure. VOICE. Dawn song, from high tree limb or treetop, a short, warbled “tio turri, weri tweee”, second and third notes lower in pitch, repeated over and over every 3–4 seconds. Day song a loud, rich series of musical notes, like that of a Pheucticus grosbeak but with emphatic upslurred note at end and with longer pauses between songs. A countersong, perhaps by female, a sliding whistle followed by two softer short low whistles. Calls include soft, high “chewp”, singly or doubled. Habitat. Variety of wooded to semi-open habitats, including Araucaria forest, dense low forest (monte), patches of shrubbery around marshes, riverine woodland (especially Celtis in Argentine coastal areas), and trees and bushes in parks and gardens. Restricted to mountainous regions in Rio de Janeiro, where most numerous at uppermost elevations, and to tree-line at 2400 m on Mt Itatiaia; otherwise mainly in lowlands.

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Food and Feeding. Much fruit, also flowers, buds and some insects; consumes sweet secretions of plant lice. In Rio Grande do Sul (Brazil) reported as eating nine species of fruit, including figs (Ficus), Japanese persimmons (Diospyros kaki) and papaya (Carica papaya). One stomach contained moyé berries (probably Schinus species). Found in pairs and in small groups, individuals often perched rather prominently in the open atop shrubbery, but quick to dive into cover when alarmed. In wooded areas regularly occurs with mixed-species flocks and will forage at almost any height, from low down to fairly high in vegetation. In Brazil appeared to glean from leaf surfaces. Breeding. Breeding reported in Oct in Rio Grande do Sul (Brazil). Builds a shallow nest 3–4 m up in tree or shrubs. Clutch 4 eggs, white or bluish-white, spotted deep red or purplish-grey, some eggs having black scrawls. No other information. Movements. Reported as rather sedentary. Possibly migratory in S Uruguay and Argentina. Status and Conservation. Not globally threatened. Fairly common in SE Brazil (from Itatiaia National Park S to Rio Grande do Sul); fairly common in Uruguay; decidedly local within most of its Argentine range. In coastal mountains of SE Brazil, where deforestation extensive, this species is largely confined to the numerous parks and reserves, among them Itatiaia, Serra dos Órgãos and Aparados da Serra National Parks. In Argentina found in Iguazú, El Palmar and Mar del Plata National Parks, as well as locally in gardens and small reserves along coast to S of city of Buenos Aires. Bibliography. Aplin (1894), Belton (1985), Burns & Racicot (2009), Descourtilz (1852), Di Giacomo (2009), Diamond & Lovejoy (1985), Frisch & Frisch (1964), Holt (1928), Hudson (1870), Isler & Isler (1999), Meyer de Schauensee (1966), Narosky & Yzurieta (1987), Ogilvie-Grant (1912), Parker & Goerck (1997), de la Peña & Rumboll (1998), Ridgely & Tudor (1989, 2009), Santos (1948), Sclater & Hudson (1888), Sedano & Burns (2010), Sick (1985, 1993), Souza (2002), Straneck (1990), Vigil (1973), Voss & Sander (1980, 1981), Wetmore (1926).

Genus IRIDOSORNIS

Lesson, 1844

96. Purplish-mantled Tanager Iridosornis porphyrocephalus French: Tangara à cape bleue

German: Purpurmanteltangare

Spanish: Tangara Capiazul

Taxonomy. Iridornis porphyrocephala P. L. Sclater, 1856, neighbourhood of Quito, Ecuador. Has been considered to form a superspecies with I. analis, and sometimes treated as conspecific; evidence for treatment as two separate species is weak, and sympatry not proven. Monotypic. Distribution. Colombia (entire W slope and very locally on E slope of W Andes, also W slope of C Andes in Antioquia) S to NW Ecuador (Carchi, Imbabura and NW Pichincha). Descriptive notes. 14 cm; three males 20·3– 23 g, one female 21·5 g. Unmistakable tanager, mostly dark blue with yellow throat, and short thick bill. Has base of forehead, lores and ocular area deep black and ear-coverts blackish, somewhat glossed dark purple, forming mask (looks blackish in most lights and does not contrast strongly with rest of plumage); crown, side of head, mantle and back intense dark purplish-blue, becoming slightly paler bluishgreen on rump (can look greenish in some lights); tail mainly dusky, rather obscurely edged and tinged dark blue; upperwing-coverts black, broadly and sharply edged dull blue, flightfeathers black, narrowly edged dull blue, tertials black, broadly edged all around with dull blue; wide throat area contrastingly bright yellow, surrounded by dusky purplish-blue that extends to breast and side of chest; central breast, sides and flanks duller and paler blue, sometimes with brownish tinge, centre of lower breast and belly buff, becoming chestnut on undertail-coverts; iris reddish-brown; upper mandible black, lower mandible bluish-grey; legs dark brownish-grey. Sexes similar. Juvenile undescribed. VOICE. Calls (in Colombia) include high, slightly raspy “tsit”, sometimes accelerating into trill. Possible song a lisping, buzzy “seeeeééu”, c. 1 second long, rising and, at end, falling. Also gives longer “ts-ts-tseééuit” tseeuwee”, which may be a song or a song variation. Habitat. Very wet mossy forest, tall second growth, and along openings or borders in forest; a few records from less humid areas. Mostly at 1500–2200 m; locally down to 750 m in Cauca and up to 2700 m in Antioquia (above Medellín), in Colombia; in Ecuador mostly c. 1200 m or somewhat higher, but few records. Food and Feeding. Berries and insects. Single individuals, pairs or families follow mixed tanager flocks or forage alone, 2–10 m up inside or at edge of forest; usually keep mostly out of sight in shrubbery. Does not gather at fruiting trees with other frugivores. Moves more slowly and forages more methodically than do Tangara tanagers, often sitting quietly for a few moments or spending 30 seconds to a minute at a location before moving. Hops and peers in dense foliage. Breeding. In Colombia, begging young with two adults in Jul in Cauca (Cerro Munchique) and nine breeding-condition birds in May–Jun at N end of W & C Andes. No other information. Movements. Presumed resident. The records at unusually high and low elevations in Cauca and Antioquia, in Colombia, may involve wandering individuals. Status and Conservation. Not globally threatened. Currently considered Near-threatened. Restricted-range species: present in Chocó EBA. Rare to locally fairly common in Colombia; very rare in Ecuador. In Colombia, may be rare or locally extirpated from portions of its range at N end of C Andes and N portion of W Andes; no recent records from Antioquia. In Ecuador, only a few scattered records in N; in addition, a specimen apparently from S Ecuador but of uncertain locality, possibly from Cañar, along Guayaquil–Quito railway. Deforestation, farming activities and illegal cultivation of coca and marijuana within this species’ range (especially in SW Colombia) are ongoing threats within the narrow elevational band on Pacific-slope montane forest where it lives. Occurs in several protected areas, including Tatamá, Farallones de Cali and Munchique National Parks and Tambito and El Pangan Nature Reserves, in Colombia; not known to occur in any reserves or protected areas in Ecuador. Bibliography. Álvarez-Rebolledo et al. (2007), Anon. (2010a), Butchart & Stattersfield (2004), Chapman (1926), David & Gosselin (2002b), Donegan & Dávalos (1999), Dunning (1982), Fjeldså & Krabbe (1990), Hellmayr (1936), Hilty & Brown (1986), Isler & Isler (1999), Meyer de Schauensee (1964, 1966, 1970a), Moore et al. (1999), Orcés (1944), Orejuela & Cantillo (1990), Remsen et al. (2010), Renjifo et al. (2002), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Sedano & Burns (2010), Stattersfield & Capper (2000).

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209

SPECIES ACCOUNTS

PLATE 10

97. Yellow-throated Tanager Iridosornis analis French: Tangara à bavette jaune

German: Gelbkehltangare

Spanish: Tangara Goliamarilla

Taxonomy. T.[anagra] analis Tschudi, 1844, Valley of Vitoc, Junín, Peru. Has been considered to form a superspecies with I. porphyrocephalus, and sometimes treated as conspecific; evidence for treatment as two separate species is weak, and sympatry not proven. Monotypic. Distribution. E slope of E Andes in S Colombia (Serranía de los Churumbelos in E Cauca–W Caquetá, and E Nariño–W Putumayo border), E slope in Ecuador (Sucumbíos S to ZamoraChinchipe) and almost entire E slope in Peru (S to Puno). Descriptive notes. 15 cm; 20–29 g. Unmistakable tanager, dark blue above, with yellow throat, and short thick bill. Plumage is mostly dull blue above, darker on crown (especially forecrown) and nape and more greenish-blue on lower back and rump; lores, ocular area and side of head black, forming inconspicuous and weakly defined small mask; upperwing-coverts blackish, edged blue, primary coverts black, flight-feathers and tertials black, edged blue (more broadly on tertials), wingtips black; tail feathers dusky, edged blue; large area on throat rich deep yellow; underparts dull tawnybuff, bluish tinge on chest, sides and flanks; undertail-coverts chestnut; iris reddish-brown; upper mandible black with bluish-grey lower edges, lower mandible bluish-grey; legs dark brownish-grey. Sexes similar. Distinguished from similar I. porphyrocephalus mainly by markedly duller appearance, especially below. Immature is like adult, but duller. VOICE. Call a thick “euu” and thinner, squeezed “tseeeer”, both dropping in pitch. Song a thin, high, somewhat buzzy “ezéétseeu”, the notes slurred together, similar to abbreviated song of Sooty Grassquit (Tiaris fuliginosus), at intervals of 10 seconds or more; song also reminiscent of that of I. porphyrocephalus. Habitat. Humid and wet montane forest, especially where thick shrubby undergrowth present; regularly along dense forest borders. At c. 1450–2000 m in Colombia (few records) and 1400– 2300 m in Ecuador; 1000–2350, rarely to 2600 m, in Peru. Food and Feeding. Insects and small fruits. Of 13 stomachs examined, three contained only vegetable matter and six only animal matter and four contained both; contents included fruit, seeds, and insects. Seen singly, in loosely associated pairs or in small family parties, which regularly follow mixed-species flocks containing other tanagers; may join feeding aggregations briefly at fruiting trees. Not so active or restless as Tangara tanagers; often tends to remain somewhat hidden in vegetation when foraging, although occasionally perches briefly on open branch. Most often observed on mossy limbs or in thick vegetation, where it maintains a rather horizontal body position when foraging. Forages from about eye level to canopy height, more often at middle levels or lower. Hops on lower branches and peers at mossy substrates, where it takes food items. Breeding. No information. Movements. Resident. Minor seasonal elevational movements believed to occur in SE Peru. Status and Conservation. Not globally threatened. Fairly common within middle-elevation montane forest; as yet unrecorded in Bolivia, although likely to be present in that country. Deforestation a cause for concern within parts of this species’ range, but it occurs in several protected areas on E slope of Andes, including Cayambe-Coca Ecological Reserve and Sangay and Podocarpus National Parks (Ecuador) and Cordillera de Colán, Tingo Maria and Manu National Parks (Peru). Present also widely in unprotected areas. Bibliography. Burns (1997a), Burns & Naoki (2004), Burns et al. (2002, 2003), Fjeldså & Krabbe (1990), Hilty & Brown (1986), Isler & Isler (1999), Lysinger et al. (2005), Mee et al. (2002), Meyer de Schauensee (1966, 1970a), Orcés (1944), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman, Donegan & Caro (2008), Salaman, Donegan & Cuervo (1999), Salaman, Stiles et al. (2002), Schulenberg (2000b), Schulenberg et al. (2007), Sedano & Burns (2010), Taczanowski (1884), Terborgh & Weske (1975), Walker (2001), Weske (1972), Zimmer (1944).

98. Golden-collared Tanager Iridosornis jelskii French: Tangara à col d’or

German: Jelskitangare

Spanish: Tangara de Jelski

Taxonomy. Iridornis Jelskii Cabanis, 1873, Maraynioc, Junín, Peru. Two subspecies recognized. Subspecies and Distribution. I. j. jelskii (Cabanis, 1873) – E slope of Peruvian Andes from E La Libertad and San Martín S to Junín. I. j. bolivianus Berlepsch, 1912 – E slope of Andes from S Peru (Cuzco) S to Bolivia (La Paz). Descriptive notes. 14 cm; 16–26 g. Distinctive colourful tanager with thick and stubby bill. Nominate race has black face and throat surrounded by broad golden-yellow area that circles rearwards from forecrown and then downwards to side of neck; crown somewhat streaked and flecked with black, nape and hindneck black; upperparts dark blue, becoming slightly paler and duller on rump; lesser and median upperwingcoverts dark blue, greater coverts dusky, edged and tipped turquoise-blue, flight-feathers blackish, narrowly edged bluish; tail blackish, tinged dark blue and edged brighter blue; black of throat bordered by band of dull greenish-chestnut on chest, this becoming rufous-chestnut on breast, belly and undertail-coverts; sides sometimes weakly smudged blue; iris reddish-brown; upper mandible black, lower mandible blue-grey; legs dusky grey. Sexes similar. Juvenile much duller than adult, with rufous wash on belly. Race bolivianus slightly smaller, forehead tinged blackish. VOICE. Soft high “seep” and “cheep” calls, sometimes two or more in a series, and thin “ti-ti-ti-ti” (of 2–5 notes), similar to those of congeners. Foraging individuals (excited) in mixed-species flock utter sharper, more forceful “tsuk!-eek” and “tsuk!-eek-ee-ee”.

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Habitat. Humid-forest borders, elfin woodland and dry scrubby hillside vegetation up to tree-line, also dense woody shrubs in páramo; sometimes found in bamboo. In most areas at c. 2900 m to 3700 m; regularly (Cuzco) down to 2200 m, occasionally to 2000 m. Food and Feeding. Of 33 stomachs examined, 24 contained only vegetable matter and four only animal matter and five contained both; contents included berries, seeds, fruit, and insects. Seen alone or in pairs, less often in family groups, individuals perching in fairly upright, alert posture as they forage or move; occurs with mixed-species flocks and also forages independently of them. Forages from below eye level to the tops of small to medium-sized trees. Hops along small branches, and leans down or flutters to take small fruits. Searches for insects along slender, bare or mosscovered branches less than 1·5 cm in diameter. Also takes some prey by gleaning or picking from foliage. Breeding. Juveniles with adult in Jun and Jul and breeding-condition birds in Nov in Huánuco (Peru). No other information. Movements. Resident; minor elevational movements may occur in response to changing food resources. Status and Conservation. Not globally threatened. Restricted-range species: present in Northeast Peruvian Cordilleras EBA and Bolivian and Peruvian Upper Yungas EBA. Uncommon to fairly common. Possibly more numerous than it appears, but inconspicuous, despite bold pattern. Deforestation, especially firewood-cutting of elfin forest and Polylepis at or near tree-line, an ongoing problem. No population trends quantified, although likely that the species has declined significantly in areas where habitat damage is severe. Occurs in a few protected areas, e.g. Tingo Maria and Manu National Parks (Peru) and Madidi National Park (Bolivia). Its distributional range includes considerable intact montane woodland and montane scrub that is unprotected, but thought to be at low risk, at least in the near future. Bibliography. David & Gosselin (2002b), Fjeldså & Krabbe (1990), Isler & Isler (1999), Meyer de Schauensee (1966, 1970a), Parker & O’Neill (1980), Remsen (1984), Ridgely & Tudor (1989, 2009), Schulenberg et al. (2007), Sedano & Burns (2010), Tallman (1974), Walker (2001), Zimmer (1944).

99. Golden-crowned Tanager Iridosornis rufivertex French: Tangara auréolé German: Goldscheiteltangare Spanish: Tangara Coronidorada Other common names: Blue-bellied Tanager (caeruleoventris) Taxonomy. Arremon rufi-vertex Lafresnaye, 1842, Bolivia; error = Bogotá region of east Andes, Colombia. May form a superspecies with I. reinhardti, and sometimes considered conspecific, although there are significant morphological differences. Race caeruleoventris may be a separate species; conversely, ignicapillus and subsimilis differ little from nominate and possibly better synonymized with latter. Four subspecies tentatively recognized. Subspecies and Distribution. I. r. caeruleoventris Chapman, 1915 – N end of W Andes (Paramillo) and N part of C Andes (Antioquia S to Tolima), in NW Colombia. I. r. ignicapillus Chapman, 1915 – SW Colombia (spottily at S end of W & C Andes). I. r. subsimilis J. T. Zimmer, 1944 – W slope of Andes in Ecuador (S to upper Chiriboga road, in Pichincha). I. r. rufivertex (Lafresnaye, 1842) – Andes of SW Venezuela (S Táchira), both slopes of adjacent E Andes of Colombia (S to Cundinamarca and on E slope in Nariño), and entire E slope of Andes in Ecuador S to N Peru (near Piura–Cajamarca border and N of Marañón Valley). Descriptive notes. 15 cm; 18–28 g. Dark but richly coloured tanager with short and thick bill. Nominate race has large circular crown patch of rich golden-orange surrounded by deep velvety black of rest of head, neck and throat; upperparts mainly deep, intense, shining purplish-blue; tail blackish, outer feathers edged dark blue; lesser upperwing-coverts deep purplish-blue (like back), median and greater coverts paler blue, flight-feathers black, inner primaries and secondaries broadly edged blue (like greater coverts), tertials black, outer half of feathers edged blue; black of throat bordered below by shining dark purplish-blue on chest, breast, belly, sides and flanks; centre of belly and undertail-coverts chestnut; iris dark reddish-brown; bill black above, pale greyish below; legs black. Sexes similar. Juvenile is dusky, has wing and tail as on adult but duller, and crown patch duller and smaller. Race subsimilis is very like nominate, but coronal patch paler and duller orange-yellow; caeruleoventris differs in having crown patch brighter orange-yellow, and vent and undertail-coverts purplish-blue; ignicapillus is similar to previous, but undertail-coverts chestnut (as on nominate). VOICE. A notably quiet species. Calls include high, thin “seeep”, also short “tsit” notes. Habitat. Dense montane forest, mossy elfin woodland and patches of stunted mossy shrubbery at or slightly above normal tree-line. At c. 2600 m to 3400 m (occasional records down to 2300 m and up to 3800 m) in Colombia; mainly 2500–3300 m (locally 2100–3550 m) in Ecuador; mostly 2900–3700 m (locally down to 2500 m) in Peru. Food and Feeding. Of three stomachs examined, one contained only vegetable matter and one only animal matter, and one contained both; contents included seeds and insects. Found generally in pairs or in small groups of up to about four individuals, most often associated with mixedspecies flocks. Forages usually c. 0·5–5 m up; tends to remain out of sight as it hops and peers in dense foliage, pops out into open briefly, or flies low across small gaps and dives quickly into cover. Breeding. In Colombia, juveniles in May in Cundinamarca and Cauca, begging juvenile in Jul in Puracé National Park, and juvenile with adult in May in Nariño, and 12 breeding-condition birds in Feb–Aug in C & N end of E Andes; in Ecuador, juveniles in Mar and Jul in NW and juvenile with adult in Aug in N; in Peru, juveniles in Jun in N Cajamarca, Sept in La Libertad and Oct in Amazonas. No other information. Movements. Apparently resident. Status and Conservation. Not globally threatened. Uncommon to fairly common throughout range. As with most upper-montane and tree-line species, firewood-cutting, land clearance and human settlement have reduced and fragmented populations to varying degrees; this species appears not, however, to be facing immediate threats. Occurs in a number of protected areas, including Paramillo, Farallones de Cali and Puracé National Parks (Colombia), Uangantes, Sangay, Cajas and Podocarpus National Parks (Ecuador) and Cordillera de Colán Reserved Zone (Peru).

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SPECIES ACCOUNTS

Bibliography. Bangs & Penard (1919), Best et al. (1993), David & Gosselin (2002b), Donegan & Dávalos (1999), Dunning (1982), Fjeldså & Krabbe (1990), Goodfellow (1901), Hilty & Brown (1983, 1986), Isler & Isler (1999), Krabbe et al. (2001), Lloyd (2009), Mazar Barnett et al. (2006), Meyer de Schauensee (1966, 1970a), Moynihan (1979), Olivares (1969), Parker et al. (1985), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman, Donegan & Caro (2008), Salaman, Donegan & Cuervo (1999), Sedano & Burns (2010), Sibley & Monroe (1990), Zimmer (1944).

100. Yellow-scarfed Tanager Iridosornis reinhardti French: Tangara de Reinhardt

German: Goldbandtangare

Spanish: Tangara de Reinhardt

Taxonomy. Iridornis reinhardti P. L. Sclater, 1865, no locality = east slope of Peruvian Cordilleras. May form a superspecies with I. rufivertex, and sometimes considered conspecific, although there are significant morphological differences. Monotypic. Distribution. E slope of Andes S of Marañón Valley (from Amazonas to Ayacucho) and Cordillera Vilcabamba (in Cuzco), in Peru. Descriptive notes. 14 cm; 19·5–28 g. Distinctive dark tanager with yellow band over head; bill stubby and thick. Has head, neck, mantle and throat black, broad golden band extending across hindcrown and downwards on each side of head to ear-coverts (resembling earmuffs or scarf across rear of head); back, rump, uppertail-coverts and breast and lower underparts, including undertail-coverts, intense purplish-blue, occasionally a small amount of chestnut in undertail-coverts; upperwing-coverts, flight-feathers and tertials black, broadly edged greenish-blue (edging not in strong contrast to rest of plumage), tail dark blue; iris dark reddish brown; upper mandible blackish, lower mandible pale grey with black tip; legs blackish. Sexes similar. Juvenile apparently undescribed. VOICE. Call a sharp, high-pitched “tzit”; also “tzizi” and “tip” and “ti’. A possible song described as a clear (high-pitched) “tsee tsee-dee”. Habitat. Humid montane forest and forest borders, elfin woodland at tree-line, and scrubby low vegetation on hillsides. At 2050–3500 m; most records at 2600–3050 m. Food and Feeding. Mostly berries and small fruits, supplemented by a few insects. Of ten stomachs examined, six contained only vegetable matter and one only animal matter, and three held both; contents included fruit, seeds and insects. Found in pairs or, more often, parties of 3–5 individuals, and about equally with mixed-species flocks and away from them. Forages from near ground to tops of smaller trees, and occasionally hops on ground; in general inconspicuous, keeps mostly within cover or appears only briefly before retreating again. Obtains food items by hopping quickly through interior of trees, especially on mossy branches, where it probes and picks at moss, including hanging tufts of moss, and sometimes drops down to ground to inspect moss-covered surfaces of roots, rocks, etc. May hang downwards to glean prey from small leaves or bare branches; has been seen moving upwards or downwards in trees rapidly, in a series of short rapid hops. Breeding. Juvenile with adult in Jun in Huánuco. No other information. Movements. Presumed resident. Status and Conservation. Not globally threatened. Restricted-range species: present in Northeast Peruvian Cordilleras EBA and Bolivian and Peruvian Upper Yungas EBA. Fairly common within narrow band of high-elevation forest which its inhabits. Forest destruction, firewood-cutting and other human activities affect the species locally, but long-term populations trends unknown. Occurs in several protected areas, including Cordillera Azul and Tingo Maria National Parks. Bibliography. Clements & Shany (2001), Fjeldså & Krabbe (1990), Isler & Isler (1999), Meyer de Schauensee (1966, 1970a), Parker & O’Neill (1980), Ridgely & Tudor (1989, 2009), Schulenberg et al. (2007), Sedano & Burns (2010), Sibley & Monroe (1990), Terborgh (1971), Vuilleumier (1970), Walker (2001), Weske (1972).

Genus PIPRAEIDEA

Swainson, 1827

101. Fawn-breasted Tanager German: Schwarzrückentangare

Bibliography. Álvarez-Rebolledo et al. (2003), Anon. (2000), Belton (1985), Best et al. (1996), Brown & Neto (1976), Burns & Naoki (2004), Burns et al. (2002, 2003), Collins & Watson (1983), Contreras & Contreras (1993), Donegan & Dávalos (1999), Dunning (1982), Fjeldså & Krabbe (1990), Goodfellow (1901), Hilty (1977, 1997, 1994, 2003), Hilty & Brown (1986), Isler & Isler (1999), Koepcke (1970), Krabbe et al. (2001), Meyer de Schauensee (1966, 1970a), Miller (1963), Mitchell (1957), Moore et al. (1999), Narosky & Di Giacomo (1993), Olivares (1969), Parker & Goerck (1997), Remsen et al. (1987), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Schulenberg et al. (2007), Sedano & Burns (2010), Sick (1985, 1993), Sick & Pabst (1968), Souza (2002), Taczanowski (1884), Walker (2001), Weske (1972), Zimmer (1943a).

Genus CHLOROCHRYSA

Spanish: Tangara de Antifaz

Chlorochrysa phoenicotis French: Calliste étincelant

Taxonomy. Tanagra melanonota Vieillot, 1819, Rio de Janeiro, Brazil. Systematic position of present species has been much debated; recent DNA evidence suggests that it is properly placed in present family and is closely related, and probably sister, to Thraupis bonariensis. Taxonomic status of SE populations may be worthy of investigation. Two subspecies recognized. Subspecies and Distribution. P. m. venezuelensis (P. L. Sclater, 1857) – mountains of N, NE, SW & S Venezuela (coastal range from Carabobo E to Distrito Federal and Miranda; Sucre and Monagas; locally in Andes from NW & S Lara S to Táchira; Cerro Yaví and Cerro Taracuniña, in N & S Amazonas; probably also mountains of SE Bolívar), and W, C & E Andes of Colombia S on both slopes to Ecuador, Peru (S on W slope to Lima, on E slope throughout), Bolivia and NW Argentina (S to Tucumán and Catamarca). P. m. melanonota (Vieillot, 1819) – S & SE Brazil (locally SW Mato Grosso; and S Bahia and E Minas Gerais S to C Rio Grande do Sul), E Paraguay, NE Argentina (Misiones and N Corrientes; NE Buenos Aires S to Punta Indio) and S Uruguay. Descriptive notes. 14 cm; 18–25 g. Unusual tanager with short, wide, hirundinid-like bill with small hook, rather short legs, long wings and short tail, and sky-blue crown. Male nominate race has crown and nape medium blue, forehead, lores, ocular area and ear-coverts deep black, forming broad mask; mantle and back dull blue, lower back and rump contrasting turquoise-blue, uppertailcoverts dark blue, tail dusky blue; upperwing-coverts blackish, rather obscurely edged dark blue, primary coverts and flight-feathers dusky, narrowly edged blue, tertials blackish, more broadly edged dull dark blue; throat and entire underparts buffy cinnamon, variable, from dark buff-cinnamon to buff and to whitish-buff; iris dark red to reddish-brown; bill dusky, with lower mandible usually grey; legs slaty. Female is similar to male, but considerably duller, especially crown, which varies from dull turquoise to dull dusky blue; back tinged brownish, rump dull turquoise (duller

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Bonaparte, 1851

102. Glistening-green Tanager

Pipraeidea melanonota French: Tangara à dos noir

than male), uppertail-coverts and tail dull dusky blue, wings duller; underparts generally paler than on male, but amount of buff to buffy whitish likewise variable. Juvenile is dingy brownish-grey above and almost without pattern; immature like female, but duller. Race venezuelensis is similar to nominate, but slightly darker above, paler below, and with iris brighter red. VOICE. Infrequently heard song a series of high “see” or “sweee” notes, varying from 4 or 5 notes given slowly to a dozen or more uttered very fast like pulsating trill; does not sing regularly, but an individual may occasionally sing over and over for several minutes. In Brazil, a second song type described, more varied and possibly with mimicry. Habitat. Bushy pastures, cultivated and small semi-open areas with large trees, along forest borders and in clearings with scattered trees; mostly humid areas. Essentially a bird of forest edge and lighter woodland over most of its range. Elevational range varies geographically; in Andes mostly c. 1500– 2500 m, occasionally to 3000 m, but as low as 400 m in N Venezuela, and in Colombia down to 900 m on Pacific slope and irregularly up to 2600 m on Sabana de Bogotá; in Ecuador mostly 1000– 2800 m, but as low as 400 m on W slope in NW Azuay, and once to 3100 m; in Peru down to 700 m on Pacific slope S to Lima, mainly 1100–2900 m on E slope (a few S Peru records as low as 600 m may represent austral migrants or wandering young); in Bolivia 350–3250 m; in Argentina occurs to 500 m in Tucumán. Nominate race found from sea-level to 2050 m (on Mt Itatiaia) in SE Brazil. Food and Feeding. Of ten stomachs examined, six contained only vegetable matter and four contained vegetable and animal matter; contents included berries, fruit pulp, seeds, and larvae. Observed also to take moths and ithominid butterflies (Lepidoptera), buds and flowers. Something of a “loner”, although sometimes associates with mixed-species flocks, and may appear at fruiting trees with other species. Behaviour may recall that of both Tersina and Tangara. Seems curiously detached, flighty at times; apt to fly off some distance, flight swift and fluid, before alighting. Forages mostly in canopy or upper levels of trees when in woodlands, but at almost all levels in more open areas, especially when feeding at fruiting trees and shrubs. Often hops along large limbs and perch-gleans; occasionally sallies up and hovers briefly beneath a leaf, or sallies to air for flying insects. Has been seen to search bromeliads and undersides of branches. Breeding. Fledgling in Jul in W Venezuela (Mérida); nest-building in Mar (Huila) and birds in breeding condition in Mar (Huila) and Jul (Valle) in Colombia; adult carrying nest material in early Mar in SW Ecuador (Tambo Negro, in Loja); and juvenile or immature in Aug in C Peru (Junín). Pair in Peru observed when carrying moss to nest-site 15 m up in clump of epiphytes on tree branch at edge of clearing; nest in Brazil was 20 m up in ferns and epiphytes on horizontal branch. No other information. Movements. Nomadic or seasonal migratory movements known, but poorly documented. In N Venezuela (coastal cordillera) present mainly Jan–Jun (presumed breeding period); in Pacific Colombia, present Nov–Mar in Anchicayá Valley at 1050 m. Populations in S of range (Argentina) may undertake some seasonal movements N during coolest months. Status and Conservation. Not globally threatened. Rare to fairly common (periodically or seasonally), and widespread. Occurs in wide range of forest-border and semi-open habitats, and may be expanding its range locally with partial opening of forest in Andes, and perhaps elsewhere. No short-term risks known. Occurs in numerous protected areas over its vast range, which encompasses large amounts of suitable unprotected habitat.

German: Rotohr-Bunttangare

Spanish: Tangara Verde

Taxonomy. Calliste phoenicotis Bonaparte, 1851, Nanegal, Ecuador. Although genus traditionally placed next to Tangara in linear taxonomic sequences, genetic data, together with glistening plumage, leg morphology, and voice, indicate that the two genera are not closely related. Monotypic. Distribution. Colombia on W slope of C Andes (Antioquia) and Pacific slope of W Andes (from Risaralda) S to Ecuador (Pichincha and El Oro). Descriptive notes. 13 cm; 20–24·5 g. Small, brilliant green, strong-legged tanager with bill thinner and longer than that of Tangara. Male is glistening emerald-green above and below (black feather bases concealed), slightly darker green on back, paler on rump; small inconspicuous tuft of shining greyish-olive feathers behind eye, immediately behind this another small (more conspicuous) tuft of bright orange feathers thickened and club-like at tips; tail blackish, central feathers greenish with bright green edges, outer rectrices edged bright green; lesser and median upperwing-coverts glistening greyish-olive (usually concealed), greater coverts black (mainly inner webs), outer webs broadly edged and tipped bright green; flightfeathers blackish, basal half of outermost primaries very narrowly edged bright green, inner primaries and all secondaries much more broadly edged bright green, tertials mostly bright green, outer half

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of inner web blackish; iris dark brown; bill black; legs blackish-grey. Female is similar to male, but slightly duller (although still highly glistening). Juvenile apparently undescribed. V OICE . All vocalizations have wheezy or lispy sound. Commonest call during foraging a high, lisping and weak 3-note to 4-note “czee, czee, czee” or more buzzy “ee-see-seez-seez”. Song in Ecuador a similar series of notes, more prolonged, “czee-czee-czeet czeet, czee czee czee”, last notes slower, this series sometimes followed by extremely rapid series of ticking notes that may accelerate, then slow or end in rapid high-pitched jumble of notes. Habitat. Canopy and borders of wet mossy forest (cloudforest) and older second growth. Recorded at c. 700–2200 m in Colombia (most records above 1000 m); in Ecuador mostly 600–1700 m, occasionally to 2000 m. Food and Feeding. Small arthropods, including larvae; also fruit. In Colombian study, insect captures accounted for 62% of foraging records and included many caterpillars, and fruit comprised remaining 38% of records; Melastomataceae berries (especially Miconia) accounted for 91% of all fruit eaten. Berries usually swallowed whole. Single individuals, pairs and family groups usually forage with mixed-species flocks containing other tanagers, and regularly visit fruiting trees. In Colombia foraged from higher part of understorey upwards to canopy of small to large trees; median foraging height (63 observations) was 9 m. Notably active and acrobatic, often hangs upside-down from twigs or leaves, or stretches upwards to reach undersides of overhead leaves; also hops or runs rapidly over mossy branches and on epiphytes, and gleans from clumps of moss, bromeliads, vine tangles, hanging dead leaves and, especially, outer foliage. Often searches for insects by hopping rapidly up through palm frond leaflets. Takes small arthropods and larvae especially from undersides of leaves; unlike many species of Tangara, does not forage by peering with head downwards. Breeding. Season evidently May–Sept in Colombia, including two breeding-condition birds in Jun at N end of C Andes, nesting evidence Apr–Jun in Valle, and begging juvenile on 6th Sept in Nariño. One nest in Colombia was a small cup hollowed in moss at medium height on side of limb. No other information. Movements. Apparently resident. Status and Conservation. Not globally threatened. Restricted-range species: present in Chocó EBA. Uncommon. Confined to narrow band of pre-montane and montane wet forest on W slope of Andes in Colombia and Ecuador, an area under intense pressure from deforestation, mining, legal and illegal farming operations, and an expanding human population. Occurs in a few protected areas, most notably Tatamá, Farallones de Cali and Munchique National Parks and Tambito and El Pangan Nature Reserves (Colombia), and probably Maquipucuna Reserve (W Ecuador). Nearterm risks to the species are probably low, but the longer-term outlook is less certain. Bibliography. Burns & Naoki (2004), Chapman (1926), Donegan & Dávalos (1999), Dunning (1982), Fjeldså & Krabbe (1990), Isler & Isler (1999), Hilty (1997, 2009a), Hilty & Brown (1986), Innes (1979), Meyer de Schauensee (1964, 1966, 1970a), Moore et al. (1999), Remsen et al. (2010), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Sedano & Burns (2010).

103. Orange-eared Tanager Chlorochrysa calliparaea French: Calliste oreillard

German: Braunohr-Bunttangare

Spanish: Tangara Orejinaranja

Taxonomy. C.[allospiza] calliparaea Tschudi, 1844, “Chinchon forests”, Junín, Peru. Although genus traditionally placed next to Tangara in linear taxonomic sequences, genetic data, together with glistening plumage, leg morphology, and voice, indicate that the two genera are not closely related. Sight records of this species in W Venezuela may refer to race bourcieri or to an undescribed race. Three subspecies recognized. Subspecies and Distribution. C. c. bourcieri (Bonaparte, 1851) – S Colombia (W slope of E Andes in Cundinamarca S to head of Magdalena Valley, in Huila; E slope of E Andes S from Caquetá) S to E Ecuador and E slope in N Peru (Huánuco S to Huallaga Valley). C. c. calliparaea (Tschudi, 1844) – E slope of Andes of Peru (Pasco S to N Ayacucho). C. c. fulgentissima Chapman, 1901 – E slope of Andes of S Peru (from Cuzco) S to Bolivia (S to NW Cochabamba). Also recorded (race uncertain) on E slope of Andes in W Venezuela (on Mérida–Barinas border). Descriptive notes. 12 cm; 14·9–21·5 g. Small, strong-legged green tanager with dark throat and narrow orange rump; bill longer and thinner than that of Tangara. Male nominate race has body plumage mainly glistening emeraldgreen, feathers with black bases; small goldenorange patch on centre of crown, prominent bright orange band across lower rump; three central pairs of tail feathers dark green, sharply edged bright green, rest mainly black, narrowly edged bright green; upperwing-coverts black, edged bright green, outer half of feathers tinged green; primary coverts black, flight-feathers black, basal half of outermost primaries edged bright green, rest of primaries and all secondaries almost entirely edged bright green, tertials edged bright green and outer web of each feather greenish, inner web blackish; throat black, joining with conspicuous, triangular-shaped, small burnt-orange patch on side of neck, otherwise glistening green below, lower breast and belly intense glistening dark blue-green; iris dark brown; bill black; legs dark grey. Female is like male, but somewhat duller green, with less bluish on belly, smaller neck patch. Immature is mainly dull green. Race bourcieri is similar to nominate, but blue of lower underparts less dark and less intense, female much duller, with grey throat, smaller orange neck patch, only hint of blue on belly; fulgentissima has crown patch yellowish, neck patch bright red, and throat and entire central area of breast and belly deep violet-blue, female extensively violetblue on throat and breast, like male but duller, retains red neck patch. VOICE. Dawn song a slow series of 4–8 or so notes, “pseet pseet pseet pseet pseet”, rather high-pitched, wheezy and lisping. Song, in Peru, described as a complex, long series of high, wiry warbles, with certain phrases repeated many times. Call a high, wheezy “seeep”. Habitat. Favours humid and wet middle-elevation mossy forest (cloudforest), forest borders, and taller second growth. At 900–2000 m, but most records c. 1000–1700 m; 1600–1800 m in upper Magdalena Valley, in Colombia. Reported in Venezuela at c. 1450 m. Food and Feeding. Small arthropods; also fruit, especially small berries, such as those of Miconia and other melastomes. Of nine stomachs examined, seven contained only vegetable matter, one only

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animal matter and one both; contents included fruit, seeds, and six caterpillars (five larger than 20 mm). Usually found singly or in pairs, and usually foraging with mixed-species flocks containing other tanagers, especially Tangara. Forages mostly between middle heights and canopy. Moves in speedy fashion, running out along mossy branches and into terminal foliage, often hanging upsidedown from twigs, petioles and leaves; spending more time searching for small arthropod prey than seeking fruit. Breeding. No information. Movements. Apparently resident. Status and Conservation. Not globally threatened. Uncommon in N of range, locally more numerous towards S. Inhabits narrow band of wet E slope pre-montane and montane forest that is under pressure in many areas from human settlement and deforestation. Occurs in some protected areas, among them Cordillera Los Picachos National Park (Colombia), Cayambe-Coca Ecological Reserve and Sangay and Podocarpus National Parks (Ecuador), Manu National Park (Peru) and Madidi National Park (Bolivia). Within its range there also is extensive intact habitat that is unprotected, but secure in at the short term. Bibliography. Burns (1997a), Burns & Naoki (2004), Burns et al. (2002, 2003), Goodfellow (1901), Hilty (1999, 2003), Hilty & Brown (1986), Isler & Isler (1999), Lysinger et al. (2005), Mee et al. (2002), Meyer de Schauensee (1964, 1966, 1970a), Remsen et al. (2010), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman, Donegan & Caro (2008), Salaman, Donegan & Cuervo (1999), Schulenberg et al. (2007), Sedano & Burns (2010), Terborgh & Weske (1975), Walker (2001), Zimmer (1930, 1943a).

104. Multicolored Tanager Chlorochrysa nitidissima French: Calliste multicolore

German: Schwarzohr-Bunttangare

Spanish: Tangara Multicolor

Taxonomy. Chlorochrysa nitidissima P. L. Sclater, 1874, Antioquia, Colombia. Although genus traditionally placed next to Tangara in linear taxonomic sequences, genetic data, together with glistening plumage, leg morphology, and voice, indicate that the two genera are not closely related. Monotypic. Distribution. Colombia: locally on both slopes of W Andes from Risaralda and extreme S Antioquia S to W Cauca (Munchique area); and C Andes at N end (Antioquia), on W slope (Quindío, and reported from Ucumari Regional Park, in Risaralda) and on E slope (Caldas). Descriptive notes. 12 cm; 17·3–21·6 g. Small, strong-legged tanager with mostly yellow head and glistening harlequin-patterned plumage; bill longer and thinner than that of Tangara. Male has forehead to mid-crown, side of head to behind eye, and throat bright golden-yellow, centre of throat golden-orange, oval black patch on ear-coverts bordered behind and below by chestnut; rear crown and nape glistening golden-green; large, triangular glistening lemon-yellow patch on mantle and upper back, lower back to uppertail-coverts glistening turquoise-blue; tail dusky, central feathers tinged green and edged bright green, outer feathers mainly edged bright green; upperwing-coverts dusky, prominently edged and tipped bright green; flight-feathers dusky, basal half of outermost primaries and entire length of rest of primaries edged bright green, secondaries and tertials with outer half of each feather green and also edged brighter green; breast and lower underparts mostly glistening turquoise-blue, sides and flanks mixed glistening green and blue, centre of lower breast and belly black, undertail-coverts glistening blue-green; iris dark brown; bill black; legs dark grey. Female is similar to male, but considerably duller, head mostly dull yellow with blackish-chestnut ear patch, golden-yellow of throat somewhat stained brownish along sides, entire mantle and back bright greenish (no yellow patch on mantle), rump glistening bluish-green, tail and wings as on male but slightly duller green; glistening green below, somewhat duller on lower underparts. Immature is like female, but even duller. VOICE. Call one to several wheezy “ceeet” or “swee” notes, similar to those of C. calliparaea. Possible song may consist of a brisk series of c. 4 wheezy “swee-see-see-see” notes. Habitat. Humid and wet mossy forest, tall older second-growth woodland and forested borders, at c. 1300–2200 m in C Andes and E slope of W Andes; very locally on Pacific slope down to 900 m. Food and Feeding. Few data. Reported food items include insects, mainly taken when foraging with mixed-species flocks in forest interior. Also fruit. Breeding. Stub-tailed juvenile with two adults on 1st Nov and grown juvenile on 8th Jan in W Andes above Cali (Valle del Cauca). No other information. Movements. Evidently resident. Status and Conservation. VULNERABLE. Restricted-range species: present in Colombian InterAndean Slopes EBA and Chocó EBA. Uncommon, local and declining. Since 1951 recorded at only two sites in C Andes, Ucumarí Regional Park (Risaralda) in early 1990s and, farther N, near Anorí (Antioquia) in 1999; in W Andes, almost all modern records are from localities in Valle del Cauca where access by ornithologists is possible. This species was formerly common, but has declined throughout range and is now recorded only infrequently and very locally, primarily in remnant forest fragments. Perhaps greatest number of recent sightings in W Andes in vicinity of Cali and N of there, including Pichindé Valley (unprotected), private forests near El 18 (Bosque de San Antonio, above Cali), Yotoco Forest Reserve, Tambito Nature Reserve and Farallones de Cali, as well as to S in Munchique National Park; also in C Andes at Otún-Quimbaya Sanctuary, in Ucumari section of Los Nevados National Park (Risaralda), and in Reserva Arrierito Antioquena (Piha Reserve) and adjacent Anorí area (N Antioquia). Other sites include Cañon del Río Barbas y Bremen (Risaralda), La Finca la Betulia–Reserva la Patasola (Quindío), La Forzosa–Santa-Gertrudis, and La Victoria. A few large tracts of intact habitat remain within its range (Farallones de Cali and Los Nevados National Parks; and Caramanta massif, near Cerro Tatamá, including Alto de Pisones area), but deforestation has been severe and continues. Furthermore, within its range an expanding network of roads is opening up ever more remote areas to logging, mining, agriculture and human occupation, all of which lead to more deforestation. This species’ range is now highly fragmented, and populations are in critical need of baseline monitoring data. Bibliography. Álvarez-Rebolledo et al. (2007), Anon. (2010a), Butchart & Stattersfield (2004), Chapman (1917), Donegan & Dávalos (1999), Dunning (1982), Hilty (1985), Hilty & Brown (1986), Isler & Isler (1999), Meyer de Schauensee (1964, 1966, 1970a), Miller (1963), Remsen et al. (2010), Renjifo et al. (2002), Ridgely & Tudor (1989), Salaman et al. (2008), Sedano & Burns (2010), Stattersfield & Capper (2000), Wege & Long (1995).

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ssp rava ssp mexicana

ssp inornata

ssp boliviana

106 ssp brasiliensis

105

ssp vieilloti

109

ssp chilensis

108

107

ssp paradisea

111 112

110

ssp cearensis

ssp cyanocephala

113

114 PLATE 11

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Family THRAUPIDAE (TANAGERS)

213

SPECIES ACCOUNTS

Genus TANGARA

PLATE 11 from limited forest disturbance. Status of brasiliensis more problematic, because of extreme deforestation in coastal region of SE Brazil; may be largely confined to protected areas, such as Sooretama and Poço das Antas Biological Reserves, among others.

Brisson, 1760

105. Turquoise Tanager Tangara mexicana French: Calliste diable-enrhumé German: Türkistangare Other common names: White-bellied Tanager (brasiliensis)

Spanish: Tangara Turquesa

Taxonomy. Tanagra mexicana Linnaeus, 1766, Cayenne, French Guiana. Based on genetic data this species is sister to T. inornata. Race brasiliensis, long regarded as a separate species because of its larger size, paler plumage and geographical isolation, has more recently been treated as a race of present species, but status merits further consideration. Proposed race lateralis (described from Apacy, on R Tapajós) is now generally considered an intergrade between nominate and boliviana; further study required. Five subspecies currently recognized. Subspecies and Distribution. T. m. vieilloti (P. L. Sclater, 1857) – Trinidad. T. m. media (Berlepsch & E. J. O. Hartert, 1902) – E Colombia (Vichada) and NE & S Venezuela (Monagas, Sucre Delta Amacuro and, S of R Orinoco, Amazonas and Bolívar). T. m. mexicana (Linnaeus, 1766) – the Guianas and Brazil (E to E Pará and S to C Mato Grosso). T. m. boliviana (Bonaparte, 1851) – E Colombia (from W Meta and R Guaviare) E through Amazonian Brazil to Pará, W Goiás and C Mato Grosso, S through E Ecuador and E Peru to N & NE Bolivia (Santa Cruz). T. m. brasiliensis (Linnaeus, 1766) – coastal SE Brazil (S Bahia S to Rio de Janeiro). Descriptive notes. 14 cm, 26 g (brasiliensis); 12 cm, 17–23·5 g (other races). Small tanager with dark, mottled appearance and contrasting yellow belly. Nominate race has mid-crown to nape and most of upperparts deep velvety black, glistening pale blue rump and uppertail-coverts somewhat mottled with black; tail black, all feathers except central pair edged dark blue; small black area encircling bill, narrow black orbital ring, faint blue supercilium, otherwise forecrown, side of head and throat to upper breast glistening cobalt-blue to purplish-blue, some black feather bases showing as mottling and spangling (especially on lower throat, and continuing to side of neck and to central breast); lesser upperwingcoverts turquoise-blue, median coverts black, broadly tipped shining blue, greater coverts black, narrowly edged blue, primary coverts black, edged blue; flight-feathers black, basal half of outer feathers edged pale blue; lower breast, belly and undertail-coverts pale yellow, sides and flanks blue to purplish-blue with heavy black scalloping and spotting; iris dark brown; bill and legs black. Sexes similar. Juvenile is mainly dull dusky blue above and dull pale yellowish-white below, with dusky on chest; immature plumage pattern more like that of adult, but duller above and below. Race boliviana differs from nominate in having lower breast to undertail-coverts richercoloured, bright creamy yellow; media differs in more well-defined blue eyebrow (but still inconspicuous), sides and flanks heavily spotted, and with little violet-blue; vieilloti has blue of head and breast darker than previous, and yellow lower underparts richer and brighter; brasiliensis is significantly larger, overall paler, more silvery blue, including lesser wing-coverts, and with white (not yellow) lower underparts. VOICE. Calls mainly rapid streams of high, chipping “tic” notes, often accelerating into trills, especially as group takes flight; higher-pitched than calls of many congeners. No song yet described. Habitat. Humid forest borders, broken second growth, lighter or disturbed forest, trees in clearings, along river and oxbow-lake edges, and along gallery forests in partly open areas; small numbers also in upper canopy and emergent treetops of lowland forest. Lowlands to c. 500 m over most of range; to 1000 m in Colombia and Venezuela (once to 2000 m on Cerro de la Neblina, in S Venezuela); isolated race brasiliensis to 500 m in SE Brazil. Food and Feeding. Arthropods and fruits. Of four stomachs examined, three contained only vegetable matter and one only animal matter; additional contents included mistletoe berries (Loranthaceae). In Trinidad, recorded as eating 26 species of fruit, of which 31% were berries of Miconia species; other fruit prominent in diet included Cecropia (13%), Ficus (10%) and Ilex (8%); in SE Brazil (race brasiliensis), recorded as taking buds of Leguminosae, melastome berries, pieces of guava fruit (Psidium), and bananas. Notably social, lives in groups of 3–10 individuals, sometimes more; usually independent of mixed-species flocks, or associates with them irregularly or for only short periods of time. In SE Brazil, more often seen in pairs. Forages from canopy down to about eye level, higher when searching for insects. In Trinidad, insect-seeking accounted for 47% of 433 foraging records, fruit for 53%; typically, took berries while perched and either swallowed them whole or mashed them; pecked pieces from larger fruit. Most insects sought on underside of rather open bare and often dead branches of c. 1·3 cm or less in diameter (91% of 201 insect-hunting records); also spent some time in searching foliage, flowerheads and seedheads, and made sallies to air. Elsewhere in range, foraging behaviour similar to that observed in Trinidad, this species favouring middle levels to high open, bare branches as foraging sites. Breeding. In E Colombia nests reported in Feb and Oct; season Apr–Oct in Trinidad and Feb–Apr and Aug in Suriname; recorded Nov in Pará, in Brazil. Pairs separate from flock to breed, but start rejoining flock intermittently as soon as eggs hatch. Co-operative breeding recorded at two nests in Trinidad, where nestlings fed by and attended by all members of group from hatching until independence. Nest built by female, attended by male, a deep cup of moss and fibres, decorated on outside with lichens and leaves, two Trinidad nests 6 m and 7·5 m above ground, one in upright fork of mango tree; reported as usually nesting high in Suriname; a Brazil nest was in branch fork in shrub. Clutch 2–3 eggs, greyish to greenish with brown or purple markings; in captivity, incubation period 12–14 days. No other information. Movements. Apparently resident. Status and Conservation. Not globally threatened. Widespread, and uncommon to locally common; small isolated population along coastal SE Brazil (brasiliensis). Found in many protected areas and also widely in unprotected areas. Primarily an “edge” species that derives some benefit

Bibliography. Beebe (1909), Belcher & Smooker (1937), Burns & Naoki (2004), Boesman (1998), Collins & Araya (1998), Descourtilz (1852), Diamond & Lovejoy (1985), Dick et al. (1984), Dunning (1982), ffrench (1991), Haverschmidt (1948, 1968), Haverschmidt & Mees (1994), Hellmayr (1910), Hilty (2003), Hilty & Brown (1986), Ingels (1974c), Isler & Isler (1999), Marantz & Zimmer (2006), Mee et al. (2002), Munn (1985), Novaes (1973), Oren & Parker (1997), Parker & Goerck (1997), Pearson (1971, 1975b), Pelzeln (1869), Pinto (1944b), Remsen et al. (2010), Restall et al. (2006), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman et al. (2008), Schulenberg et al. (2007), Sedano & Burns (2010), Sick (1985, 1993), Snow, B.K. & Snow (1971), Snow, D.W. & Collins (1962), Snyder (1966), Souza (2002), Sproule (2006), Terborgh & Weske (1969), Tostain et al. (1992), Willis (1977), Yuri & Mindell (2002), Zimmer (1943c).

106. Plain-colored Tanager Tangara inornata French: Calliste gris German: Schlichttangare Other common names: Plain Tanager

Spanish: Tangara Cenicienta

Taxonomy. Calliste inornata Gould, 1855, Santa Fé de Bogotá, Colombia. Based on genetic data this species is sister to T. mexicana. Three subspecies recognized. Subspecies and Distribution. T. i. rava Wetmore, 1963 – Caribbean slope in Costa Rica (S from Heredia) and Panama (S to Colón and Coclé). T. i. languens Bangs & Barbour, 1922 – Panama (Caribbean side E from Colón and Coclé, and Pacific slope E from W Panamá) E to Colombia (Chocó, and S in Pacific lowlands to N Valle del Cauca). T. i. inornata (Gould, 1855) – N Colombia at N base of W Andes (Sinú Valley in Antioquia), lower Cauca Valley (in Antioquia), and middle Magdalena Valley (from Antioquia S to Caldas, Santander, Boyacá and Cundinamarca). Descriptive notes. 12 cm; 16·4–19·1 g. One of the few members of genus that lacks colourful plumage. Nominate race is mainly dark lead-grey on head and back, slightly paler grey on rump and uppertail-coverts; lesser upperwing-coverts bright blue (usually concealed), rest of coverts, flight-feathers and tail plain blackish; throat, chest and sides plain neutralgrey, fading to dull whitish (or faintly tinged buff-white) on centre of breast, belly and undertail-coverts; underwing-coverts white; iris dark brown; bill and legs black. Sexes similar. Immature is duller and even plainer than adult. Race rava is similar to nominate, but has faint buff tinge on breast, more prominent buff on belly and undertail-coverts; languens differs in having head and upperparts decidedly paler, more plain grey, rump slightly paler than back, and throat and chest slightly paler grey than nominate, central breast tinged buffy white (not white). VOICE. Song of c. 15 notes, a short fast series of “tsst” that gradually slows; also described as a random series of high thin whistles, short staccato notes and sputters, “tseee tsp tsp tsee tsp tseee tseee tsp tsp tsp”. Calls mostly high-pitched, sibilant, and not especially distinctive, including “tst”, “tsit”, “tseep” and “jeet”, singly or sometimes accelerating into rapid dry trill; short “trrr” in flight. Habitat. Humid forest borders, second growth and disturbed or semi-open areas, gardens, parks, plantations and urban areas with trees; occasionally forest canopy. Lowlands to c. 400 m in foothills in Costa Rica; to 800 m in Panama and to 1000 m in Colombia. Food and Feeding. Arthropods and fruit. In Panama recorded as eating wide variety of small (3–5 mm) fruits, including those of Tetracera (Dilleniaceae), Lantana (Verbenaceae) and Hamelia (Rubiaceae), and catkins of Cecropia; last-mentioned commonly consumed throughout range. Lives in pairs or in groups of 3–4 when breeding; in groups of up to about six individuals, occasionally larger groups, outside breeding season. Nervous, fidgety, almost hyperactive, and often noisy as it constantly flicks wings and tail, and flies from tree to tree. Forages mainly in canopy of trees, but descends to shrubs for fruit. Breeding. All information from Panama. Season Mar–Aug. At one nest two nestlings fed by four adults, which visited together. Nest a tidy cup of light-coloured fibres covered with green moss, spider webs and cocoon silk; each of three nests c. 9 m up in foliage of fruiting tree, another nest 20 m up. Clutch 2 eggs, whitish, speckled. No other information. Movements. Resident. Reported seasonal variation in numbers in C Panama, suggesting some regional movements. Status and Conservation. Not globally threatened. Uncommon to fairly common in much of range. Found in several protected areas, including Metropolitan, Soberanía, Chagres and Darién National Parks (Panama) and Los Katíos National Park and El Paujíl Reserve (Colombia). This species’ range also includes much suitable second-growth and forest-edge habitat. Unlikely to face any risks in immediate future. Bibliography. Burns & Naoki (2004), Davis (1972), Dunning (1982), Eisenmann (1952), Garrigues & Dean (2007), Greenberg (1981c), Hilty & Brown (1986), Isler & Isler (1999), Karr (1977), Leck (1971a, 1971b, 1972c, 1975), Meyer de Schauensee (1964, 1966, 1970a), Moynihan (1962c), Olivares (1969), Remsen et al. (2010), Ricklefs (1976), Ridgely & Gwynne (1989), Salaman et al. (2008), Sedano & Burns (2010), Skutch (1954, 1976), Slud (1964), Stiles (1983), Stiles & Skutch (1989), Strauch (1977), Wetmore et al. (1984), Willis (1980), Willis & Eisenmann (1979).

107. Azure-rumped Tanager Tangara cabanisi French: Calliste azuré German: Cabanistangare Other common names: Cabanis’s Tanager

Spanish: Tangara Chiapaneca

Taxonomy. Calliste cabanisi P. L. Sclater, 1868, Costa Cuca, Guatemala.

On following pages: 108. Grey-and-gold Tanager (Tangara palmeri); 109. Paradise Tanager (Tangara chilensis); 110. Seven-colored Tanager (Tangara fastuosa); 111. Green-headed Tanager (Tangara seledon); 112. Red-necked Tanager (Tangara cyanocephala); 113. Brassy-breasted Tanager (Tangara desmaresti); 114. Gilt-edged Tanager (Tangara cyanoventris).

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Family THRAUPIDAE (TANAGERS)

PLATE 11

SPECIES ACCOUNTS

Originally described under name Calliste sclateri, but that name is invalid, as preoccupied. On basis of plumage, habitat and voice appears most closely related to T. palmeri. Monotypic. Distribution. S Mexico (Sierra Madre de Chiapas) and adjacent SW Guatemala. Descriptive notes. 14 cm. Pale blue-grey tanager with black-speckled collar across whitish chest. Has crown bluish-grey, becoming silvery blue on nape, bluish-green on mantle and azure-blue on rump; crown and back somewhat speckled and scaled with dusky; lesser upperwing-coverts mostly pale blue (turquoise-blue), median and greater coverts black, edged bluish, flight-feathers black, all except outermost primaries edged pale blue, tertials black, more broadly edged pale blue; tail feathers blackish, narrowly edged pale blue; lores black, black extending narrowly over bill, and blackish line across lower ear-coverts (variable in extent and length); otherwise, face and entire underparts pale bluish-white to greenish-white (varies with light), lower throat, chest and upper breast somewhat scaled and mottled with black (forming speckled collar); iris dark brown; bill dark grey, dusky tip; legs blackish. Differs from similar T. palmeri mainly in smaller size and more bluish appearance. Sexes similar. Juvenile is like adult, but duller, crown greyish with pale blue tinge, back similar but with small amount of dark mottling, lesser wing-coverts greyish, flight-feathers edged duller blue, lores blackish, lower edge of ear-coverts with variable number of dark markings, whitish below, breast with minor dark mottling (amount variable), bill grey; older juvenile soon moults into plumage essentially identical to that of adult. VOICE. Calls include high, rising “seee”, rapid “chi-di-di-dit”, high tittering notes and, in chases in flight, a buzzier “bzzz bzzz bzzz zeeah” and other notes. Song, delivered from high bare treetop branch, apparently a high, thin “seeeeawit, chi-di-di-di”, sometimes repeated in rapid succession; may sing for several minutes from one perch, then move to another and continue singing. Habitat. Tall humid broadleaf evergreen forest, forest borders and small plantation-like clearings with scattered trees, at 600–1800 m; most records above 1000 m, mainly 1250–1650 m. Food and Feeding. Observed to eat figs (Ficus), melastome berries and insects in Mexico. Most often seen singly or in pairs, but noted also in small groups, and once a party of 26 individuals; occasionally forages with mixed-species flocks, but apparently more often independent of them. Active and noisy bouts of foraging interrupted by periods of quiet and rest in tops of large trees. Forages mostly in upper levels and canopy of forest, but sometimes lower, and once on ground at base of melastome shrub. Insect-searching behaviour included peering at leaves and leaf axils, examining undersides of outer branches by hopping and leaning down, first on one side, then on other, to inspect mossy branches; also examined mossy branches less than 5 cm in diameter, and often executed short clumsy or heavy aerial sallies from exposed perches above canopy. Breeding. Breeding recorded in early wet season, mid-Apr to mid-Jun, in Mexico although wet season extends to Sept. Co-operative breeding with helpers recorded; 4–6 birds in adult plumage observed at nest, although not known if all contributed food. Most of following details from a yearlong study in which eight nests were located. Nest material brought by both adults, but more of construction work carried out by one member of pair (presumably female); cup-nest of moss and lichen, placed usually in fork where foliage densest and near end of long horizontal branch in upper half of large tree (taller than 15 m), five of eight nests in Ficus cookii; also, one nest reported in pine (Pinus), one in exotic Cupressus, another in an Inga growing over coffee, and one mossy cup-nest was 12 m above ground in branch fork c. 1·5 m from end of large limb of fig tree overhanging canyon. Clutch apparently 2 eggs (inferred from number of nestlings), undescribed; incubation presumably by female alone, at two nests for, respectively, 73·9% and 81·7% of daylight hours, average length of incubation stint 18·7 minutes (average recess 6·7 minutes) at one nest and 20·7 minutes (average recess 5·7 minutes) at other, incubation period c. 14 days; chicks fed by both adults, also by helpers when present, c. 5–8 feeding visits per hour, helpers accounted for 4% of feeds at onset and up to 38% on day before young left nest; nestling period c. 15 days. Data from six nests indicated nest success c. 29–43%. Movements. Resident. During year-round study believed to remain on territory throughout. Status and Conservation. ENDANGERED. Restricted-range species: present in North Central American Highlands EBA. Locally common to rare. An extremely localized and declining species. Known from five localities in SE Mexico, but all recent reports from El Triunfo Biosphere Reserve (mainly in buffer zone, which includes numerous expanding settlements). In adjacent Guatemala presently known from eight sites, including S slope of Volcán Santa María, Dos Marías Reserve, San Maracos, Volcán Tajumulco, Volcán Atitlán, and middle valley of R Madre Vieja N of Pochuta. Most of these sites have some legal protection, including as private nature reserves, and include coffee plantations and primary forest. Within these very restricted areas the species appears to be locally common. Expanding human settlement and loss of habitat from deforestation are main threats; some revenue from tourism and birdwatching at these sites, however, is helping local communities to realize a benefit from conserving these areas. Bibliography. Álvarez del Toro (1964), Anon. (2010a), Blake (1953), Brodkorb (1939), Butchart & Stattersfield (2004), Davis (1972), Eisenmann (1955), Eisermann, Arbeiter, López, Avendaño & de León Lux (2011), Eisermann, Arbeiter, López, Avendaño, de León Lux, Burge et al. (2011), Eisermann, López et al. (2011), Gómez de Silva et al. (1999), Heath & Long (1991), Hilty & Simon (1977), Howell & Webb (1995), Isler & Isler (1999), Land (1970), Long & Heath (1994), de Silva Garza (1997), Stattersfield & Capper (2000).

108. Grey-and-gold Tanager Tangara palmeri French: Calliste or-gris

German: Palmertangare

Spanish: Tangara Grisdorada

Taxonomy. Calospiza palmeri Hellmayr, 1909, River Sipí, Chocó, Colombia. On basis of plumage, habitat and voice appears most closely related to T. cabanisi. Monotypic. Distribution. E Panama (including Cerro Tacarcuna, Altos de Quía, Cerro Pirre and Cerro Sapo; sight records from Nusagandi, in E San Blas province, and Cerro Azul), and Pacific slope and lowlands of W Colombia and W Ecuador (S to Esmeraldas and Pichincha; also Cordillera de Chindul, in N Manabí). Descriptive notes. 15 cm; 30–32·6 g. Fairly large, plump, predominantly grey and black Tangara with black-speckled foreparts; “gold” colour inconspicuous. Plumage is mostly light pearly grey above (depending on angle of light), the grey of mantle mixed with black speckling and sprinkled with gold and silvery green, scapulars black; short black mask across base of forehead and extending through lores to just behind eye and to chin; lesser upperwing-coverts medium grey, median and greater coverts black, edged and tipped grey, flight-feathers and tertials black, edged grey; tail feathers black, edged grey; throat, side of head and median underparts off-white, variable black

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speckling of neck extending down across lower side of chest to form irregular necklace of black speckling, this mixed with gold across pectoral area; sides and flanks tinged medium grey, undertail-coverts white; iris dark brown; bill black; legs dark grey. Sexes similar. Juvenile is much duller and dingier than adult, with bill greyish and no gold on chest; soon moults into plumage that closely resembles that of adult. VOICE. Unlike most congeners, a notably vocal species. Several semi-musical calls and staccato notes, most common of which is a sweet “chup chup-sweeeeet”, last long note rising and often given over and over, or any of these notes given singly. Pair-members greet and fly off rapidly with energetic series of “chip”, “chup”, “sweet” and other notes. Habitat. Mainly humid to very wet (pluvial) forest, often on steep hillsides, forest borders, tall second growth and scattered trees in clearings. Lowlands to c. 1100 m; most numerous above c. 300 m in Colombia and above c. 400 m in Ecuador. Food and Feeding. Fruits and berries, and arthropods. In W Colombia observed to eat more than 32 species of fruit (80% of 541 observations); Miconia berries accounted for 40% of all fruit eaten, Cecropia a further 35%, and figs (Ficus) 7%. One individual seen to take insects to a nestling. Lives in pairs or in lively, excitable groups of 3–8 individuals, and may forage with mixed-species canopy flocks for short periods, but typically moves too rapidly for other species to follow and apt to fly off long distances before alighting again; reported in about half of mixed-species flocks at one site in upper Anchicayá Valley (W Valle del Cauca), in Colombia. Forages mainly in canopy or upper levels of trees, occasionally down to middle heights, rarely down to 3 m above ground; unlike congeners, does not regularly descend low to fruiting shrubs. Median foraging height in 450 observations of foraging was c. 12 m. Typically, takes berries and small fruits from upright perched position by leaning out to pluck the item, rarely plucks fruit while in flight; often takes Cecropia catkins by hanging with head downwards. Small berries swallowed whole; eats large ones piecemeal by pecking and squeezing or mashing pieces before swallowing. Almost all insects captured during aerial sallies (82% of 106 insect-hunting observations). Often perched near flowers that attracted insects, and was observed to sally to foliage for arthropod prey. May hop along treetop branches or explore foliage, but does not employ “leaning-down” search motion of many congeners. One member of a pair was seen to be struck and killed in flight by a Bat Falcon (Falco rufigularis) in Colombia. Breeding. Nests found in early Jul, early Aug and Dec; nest-building seen in Jan, and nest in Feb and pair feeding fledgling in Apr; in upper Anchicayá Valley (W Colombia) some breeding may occur all year. Nest a mossy cup placed c. 30 m above ground on large, moss-covered branch. No other information. Movements. Resident; no evidence of any seasonal movements in W Colombia. Status and Conservation. Not globally threatened. Uncommon to fairly common. Occurs at rather low density or distributed locally in Chocó–Pacific forests. Occurs in a few protected sites, such as Los Katíos National Park, possibly also Farallones de Cali National Park and El Pangan Nature Reserve, in Colombia. Species’ range includes considerable intact forest that is unprotected, but much of this (at least in Colombia) may ultimately be at risk in the long term. The species persists, in small numbers, in tall old second growth or partly disturbed areas. In Ecuador most numerous in Esmeraldas, but continues to be reported at Tinalandia, a privately held property in Pichincha; current status in Cordillera de Chindul, in Manabí, uncertain. Baseline population-monitoring desirable. Bibliography. Angehr et al. (2004), Dunning (1982), Hilty (1981, 1997, 2009a), Hilty & Brown (1986), Hilty & Simon (1977), Isler & Isler (1999), Jahn et al. (2002), Lehmann (1957), Meyer de Schauensee (1951, 1952b, 1964, 1966, 1970a), Naoki (2003c), Pearman (1993), Ridgely & Greenfield (2001a, 2001b), Ridgely & Gwynne (1989), Ridgely & Tudor (1989, 2009), Robbins et al. (1985), Salaman et al. (2008), Sedano & Burns (2010), Wetmore et al. (1984).

109. Paradise Tanager Tangara chilensis French: Calliste septicolore

German: Siebenfarbentangare

Spanish: Tangara del Paraíso

Taxonomy. Aglaïa Chilensis Vigors, 1832, no locality = Bolivia. Molecular data indicate that this species is most closely related to T. velia and T. callophrys. Race caelicolor intergrades with paradisea in E Venezuela and Guyana. Race caelicolor often misspelt “coelicolor” but former is the original spelling. Four subspecies recognized. Subspecies and Distribution. T. c. paradisea (Swainson, 1837) – SE Venezuela, the Guianas and N Brazil (S to N bank of R Amazon). T. c. caelicolor (P. L. Sclater, 1851) – E Colombia (Meta, Guainía and Vaupés), S Venezuela (Amazonas) and NW Brazil (R Uaupés and upper R Negro). T. c. chilensis (Vigors, 1832) – S & SE Colombia (W slope of E Andes in upper Magdalena Valley, in Huila; and Caquetá, Putumayo and Amazonas) S to E Ecuador, E Peru and Bolivia (S to Santa Cruz), and E through W Brazil (E to Rondônia, SE Amazonas E Mato Grosso and SE Pará). T. c. chlorocorys J. T. Zimmer, 1929 – N Peru (upper Huallaga Valley). Descriptive notes. 12–13 cm; 16–17 g (paradisea), 17–27 g (nominate and chlorocorys). Unmistakable tanager with a harlequin pattern of contrasting colors. Nominate race has most of crown and side of head scaly apple-green (feathers with black bases and pale green tips), narrow black line over bill and prominent (almost clown-like) black orbital ring; hindcrown, nape and back deep velvety black, lower back and rump entirely bright red or (e.g. Tres Esquinas area of Caquetá, in Colombia) changing to orange narrowly on uppertail-coverts; tail uniformly black; bend of wing and lesser upperwing-coverts turquoise, median coverts cobalt-blue, greater coverts, flight-feathers and tertials deep black, basal half of primaries (except innermost ones), as well as primary coverts, edged cobalt-blue to purplish-blue; throat to central chest glistening violet, breast, sides and flanks glistening turquoise-

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blue, centre of belly and undertail-coverts deep black; iris dark brown; bill jet-black; legs dark grey to black. Sexes similar. Immature is similar to adult, but duller, with lower back variously orange to orange-red. Race caelicolor is similar to nominate, but lower back and most of rump flamered and uppertail-coverts contrastingly bright golden-yellow; paradisea differs from previous mainly in having red of lower back slightly paler and more restricted in extent, uppertail-coverts lighter yellow, also differs from nominate in smaller size, and median wing-coverts same colour as throat (not paler cobalt to turquoise); chlorocorys is similar to caelicolor, but larger, green of top of head extending farther rearwards, and green colour is brighter, more yellowish-tinged. VOICE. Noisier than most congeners. Individuals, especially when in flock, almost constantly give rapid, high-pitched chipping notes, including sharp “chak”, and a rising “zeet” or “swee”, either singly or in excited series, sometimes rapidly. Notes given in flight and during foraging or moving rapidly through canopy. Dawn song “cheek, cheet” (second note may be slightly higher-pitched) repeated over and over in long, somewhat irregular series, average c. 13 notes in 10 seconds; also described, in Peru, as a regularly spaced series of “swee? tchip” phrases. Day song in SE Peru “tsip seeeawee”, repeated over and over. Habitat. Everywhere found in high canopy and treetops, and often in canopy of emergent trees (leafless or with foliage) such as Ceiba pentandra. In Amazonian lowlands, occurs in tall humid várzea and terra firme forest, somewhat less frequently in forest borders or tall second growth, or tall trees in clearings. In humid and wet Andean foothills and at lower elevations, occupies variety of forested habitats, including broken landslide and steep hillside forest, occasionally even rather scrubby woodland. Lowlands to c. 1500 m or slightly higher. Food and Feeding. Fruits and small berries, also arthropods; observed to eat Miconia berries and fruits of Aralia. Of twelve stomachs examined, ten contained only vegetable matter and one only animal matter and one contained both; contents included fruit pulp and insects. Other stomachs contained seeds, fly larvae (Diptera), short-horned grasshoppers (Caelifera) and spiders (Araneae). At one nest chicks fed with spiders, orthopterans and caterpillars. Excitable and noisy, and notably gregarious, almost always encountered in single-species flocks of 3–15 individuals, sometimes more, or in groups that associate with canopy mixed-species flocks; regularly associated with T. schrankii, and the two species may forage together, either accompanying mixed flocks or travelling independently of them. Moves in and out of slower-moving mixed-species flocks, sometimes remaining for only a few minutes before flying off, abandoning a mixed flock. Forages mainly very high up, averages 25–50 m above ground, only infrequently dropping low along forested edges to reach fruiting shrubs. Groups restless and active, individuals searching rapidly on high open branches almost exclusively by hopping outwards and leaning downwards, first along one side of a branch and then the other, to check sides and undersurface; searches both bare and mossy branches. Also searches in foliage for small berries. Breeding. In Peru, breeding recorded in Jun–Sept in SE and nest-building observed in Nov in C (Pasco); nest reported in Oct in French Guiana. Nest in SE Peru a small cup built mostly by female, both adults visited during construction but only one (presumably female) carried material, including green lichen-like moss, a fungus (Rhizomorpha corynephora), fluff from seeds like those of dandelion (Taraxacum), possibly spider web, and straw-like grass lining, placed 31 m up in small outer branch fork of Ceiba pentandra tree; no information on clutch size, incubation stints variable, 15–55 minutes, incubation period less than 16 days; nestlings fed by both parents, by which time tree had lost all its leaves and nest was exposed; nest ultimately failed following disturbance (unknown causes); c. 1 month later a second nest was under construction in same tree, disappeared after 8 days. In Brazil, cup-nest 5–20 m above ground reported; eggs apparently white or greenishwhite with lilac and purplish-red markings, especially around large end. No further information available. Movements. Resident. Status and Conservation. Not globally threatened. Widespread, and fairly common to common. Found in many protected areas, and widely distributed in unprotected areas. This species does not face any immediate threats, although it suffers adverse impact locally from destruction of Amazonian rainforest. Bibliography. Bock (1995), Burns & Naoki (2004), Burns et al. (2002, 2003), Chapman (1926), Clements & Shany (2001), Delacour (1948), Descourtilz (1852), Dick et al. (1984), Donahue & Wood (1992), Dunning (1982), Gilliard (1941), Haverschmidt & Mees (1994), Hilty (1994, 2003), Hilty & Brown (1986), Isler & Isler (1999), Marantz & Zimmer (2006), McCarthy (2006), Mee et al. (2002), Meyer de Schauensee (1951, 1952b, 1964, 1966, 1970a), Meyer de Schauensee & Phelps (1978), Moore et al. (2009), Munn (1985), Ogilvie-Grant (1912), O’Neill (1974), Ottavio et al. (2008), Pacheco et al. (2007), Parker et al. (1982), Pearson (1971), Phelps & Phelps (1963), Ridgely & Greenfield (2001a, 2001b), Ridgely & Tudor (1989, 2009), Salaman, Donegan & Caro (2008), Salaman, Donegan & Cuervo (1999), Schulenberg et al. (2007), Sedano & Burns (2010), Sick (1985, 1993), Snethlage (1914), Snyder (1966), Souza (2002), Taczanowski (1884), Terborgh & Weske (1969), Tostain et al. (1992), Willis (1977), Wood et al. (1992), Zimmer (1943b).

110. Seven-colored Tanager Tangara fastuosa French: Calliste superbe

German: Vielfarbentangare

Spanish: Tangara Sietecolores

Taxonomy. Tanagra fastuosa Lesson, 1831, Pernambuco, Brazil. Recent molecular-genetic studies suggest that this species and T. seledon are likely sister-species and form a monophyletic clade with T. cyanoventris, T. cyanocephala and T. desmaresti. Although apparently illustrated as early as 1648, not formally described until 183 years later. Monotypic. Distribution. Extreme E Brazil: Paraíba, Pernambuco and Alagoas. Descriptive notes. 13 cm. A brilliantly coloured and boldly patterned tanager, with bill relatively thick for the genus. Has narrow black ocular ring and line surrounding base of bill; otherwise, entire head to hindneck and upper throat bright greenish-turquoise, mantle and scapulars deep black, back, rump and uppertailcoverts bright burnished yellow-orange; tail feathers black, edged deep blue; lesser upperwing-coverts turquoise-blue, median coverts cobalt-blue, greater coverts black, broadly edged deep violet-blue, flight-feathers black, edged dark blue, tertials black, broadly edged orange-yellow on outer webs; lower throat black, the black extending narrowly onto side of neck to meet black of back; chest light blue, shading to deep ultramarine-blue on lower breast, belly and undertail-coverts; iris dark brown; bill black; legs dark grey. Sexes similar. Juvenile undescribed. VOICE. In disputes or when excited, a high “ti-ti-ti-ti”, much like that of many congeners. No other information.

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Habitat. Humid floodplain-forest and upland forest, forest borders and older second-growth woodland in lowlands and foothills, to c. 1000 m. Occurs also in degraded habitat composed predominantly of Cecropia and Parkia, although long-term viability in such habitats unknown. Possibly also in somewhat drier, more open habitat (cerrado); confirmation needed. Food and Feeding. Fruits and arthropods One stomach contained vegetable matter. Occurs in pairs and in small groups, often with mixed-species flocks, staying mostly in upper levels and canopy, but descending low along forest borders and in young second growth, gardens, and orchards with bromeliads. Searches for insects by hopping along branches and peering down on one side and then the other. Behaviour similar to that of T. chilensis and T. seledon. Breeding. Breeding reported in Oct–Mar (austral spring and summer). Nests reported in forest middle storey, usually in large bromeliads, which could be keystone plants in breeding requirements. No other information. Movements. Resident. Status and Conservation. VULNERABLE. Restricted-range species: present in Atlantic Slope of Alagoas and Pernambuco EBA. Uncommon to rare; declining within its very small global range. Confirmed sites (more than 50) for this species are in areas of Atlantic Forest. It has been recorded also in Rio Grande do Norte (at Capim Macio and Parque das Dunas Costeiras, and Baía Formosa), but these sites are cerrado, a significantly different habitat from those where the species has been found previously; records possibly involve escaped cagebirds. As late as 1987, the species was still relatively numerous in forests of Serra Branca, near Murici, in Alagoas. Since then it has declined steadily, as a result of severe deforestation coupled with trapping for cagebird trade. Estimated that no more than 2% of original Atlantic Forest remains in NE Brazil, the consequence of logging, and conversion to sugar-cane plantations and cattle pastures; few, if any, forest tracts larger than 4000 ha remain in this area, and sites are threatened by fires from adjacent plantations and by new roads (which facilitate logging). Intense trapping also continues, because of high price commanded by this species in trade. These factors, combined with its extremely limited range, place this species at significant risk. Known to occur in a number of protected areas, including Dunas de Natal State Park, Charles Darwin Ecological Refuge, Mata do Pau Ferro Ecological Park, Serra dos Cavalos UFPE and Tapacurá Ecological Stations, and Saltinho and Pedra Talhada Biological Reserves. Reforestation projects underway in Pedra Talhada and elsewhere may eventually increase the area of available habitat. Bibliography. Anon. (2010a), Burns & Naoki (2004), Butchart & Stattersfield (2004), Carvalho (1986), Forbes (1881), Isler & Isler (1999), Lamm (1948), Ridgely & Tudor (1989, 2009), Sedano & Burns (2010), Sick (1985, 1993), Silveira et al. (2003), Souza (2002), Stattersfield & Capper (2000).

111. Green-headed Tanager Tangara seledon French: Calliste à tête verte

German: Dreifarbentangare

Spanish: Tangara Arcoiris

Taxonomy. Tanagra Seledon Statius Müller, 1776, Cayenne; error = Rio de Janeiro, Brazil. Recent molecular-genetic studies suggest that this species and T. fastuosa are likely sister-species and form a monophyletic clade with T. cyanoventris, T. cyanocephala and T. desmaresti. Monotypic. Distribution. SE Brazil (from SE Bahia S to Santa Catarina and NE Rio Grande do Sul) and adjacent parts of SE Paraguay (Alto Paraná) and NE Argentina (Misiones). Descriptive notes. 13 cm; mean 18·7 g (nine birds from Paraguay), two others 16 g and 20 g. Brightly coloured tanager with complex plumage pattern reminiscent of that of T. chilensis and T. fastuosa, but greener and more pastel. Male has narrow black line surrounding bill and narrow black orbital ring; otherwise, entire head, nape and chin bright aquamarinegreen, broad band across nape and wrapping around on sides of neck to throat shining pale lime-green (varying to burnished yellowishgreen); back and scapulars mostly black, rump burnished yellow-orange, uppertail-coverts bright apple-green; tail black, edged pale green basally, changing to turquoise-blue distally; upperwing-coverts rich dark violet-blue, alula and primary coverts black, edged violet-blue on outer webs; flight-feathers black, broadly edged light green; lower throat and central upper chest black, which extends variably over side of chest; breast and central belly bright turquoise-blue, sides of belly, flanks and undertail-coverts bright applegreen; iris dark brown; bill black; legs dark grey to black. Female is similar to male or slightly duller. Immature is much duller than adult. VOICE. Call a high, slightly buzzy “zweet” with rising inflection, and sometimes followed by other excited notes, e.g. “zwéé-dit, zwéé-dit-see” or “zwéédi-di-di-dit”, and other twittering notes. Dawn song, from crown of tree, a high-pitched, steadily repeated, trisyllabic “sít sir zwéét”, up to 90 in a minute. Habitat. Found in canopy of humid forest, along forest borders, in small clearings with scattered trees and shrubs, and in second growth of various ages, also parks, gardens and around human dwellings. Lowlands to c. 900 m; most numerous at low elevations. Food and Feeding. Fruits, also arthropods. Takes wide variety of cultivated fruits, including oranges, papayas (Carica papaya) and bananas, as well as many wild fruits, such as figs (Ficus), Hamelia, Urtica, and berries from bromeliads; in Paraguay observed to eat fruit of Allophyllus edulis. Of six stomachs examined, five contained only vegetable matter and one only animal matter; contents included fruit and ants (Formicidae). Occurs in pairs and in small groups, sometimes in larger groups containing up to 20 individuals; pairs or groups forage alone or associate with mixed-species flocks. Forages mostly from middle heights to canopy (c. 9–25 m up) in forest, sometimes lower along forest borders and in disturbed areas. Foraging behaviour versatile; often employs acrobatic movements when hopping along branches, or peering and gleaning from leaf surfaces, bark, lichens and epiphytes, and regularly leans down to inspect sides of bare branches. In Paraguay obtained fruit of Allophyllus edulis mostly by perching and reaching for fruits, which it often manipulated with bill to remove seeds or soften pulp. Will visit feeding trays. Breeding. Season Nov–Feb in Brazil, and recorded in Nov and Dec in Paraguay and Nov in NE Argentina; apparently, second brood often attempted shortly after completion of first. Nest reportedly built by both sexes, a compact cup of grass and leaves, lined with softer material, and concealed in foliage of tree or shrub, in bromeliad along side of trunk or among epiphytic orchids on limb; nests also reported in mature banana plants. Clutch usually 3 eggs (once 2), white or pinkish-white with various shades of brown and grey markings, especially at large end; no information on incubation and nestling periods in the wild; young from first and second broods may accompany adults for most of first year. In captivity: incubation period reported as 17 days; young leave nest after 30–35 days, but fed by parents up to 75 days. Movements. Resident. Possible seasonal movements between forest and semi-open areas reported.

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Status and Conservation. Not globally threatened. Uncommon to locally fairly common. Despite occurrence in variety of disturbed and second-growth habitats, this species has, since late 1980s or earlier, disappeared from large areas of recently deforested terrain, and is also absent from remnant woodlots studied in São Paulo state of SE Brazil; this suggests that total numbers are much reduced and distribution highly fragmented. Viable populations now confined mainly to protected areas, among them Sooretama, Poço das Antas and Augusto Ruschi (Nova Lombardia) Biological Reserves and Itatiaia, Tijuca, Serra dos Órgãos and Iguaçu National Parks (Brazil) and Iguazú National Park (Argentina). Bibliography. Athanas (2010a), Belton (1974), Bertoni (1901), Burns & Naoki (2004), Descourtilz (1852), Diamond & Lovejoy (1985), Euler (1867, 1900), Goeldi (1894), Hempel (1949), Isler & Isler (1999), Mitchell (1957), Narosky & Di Giacomo (1993), Nehrkorn (1899), Norgaard-Olesen (1973), de Oliveira & Rech (1981), de la Peña & Rumboll (1998), Ridgely & Tudor (1989, 2009), Sazima et al. (1993), Sedano & Burns (2010), Sick (1985, 1993), Snethlage & Schreiner (1929), Souza (2002), Vigil (1973), Willis (1979).

112. Red-necked Tanager Tangara cyanocephala French: Calliste à tête bleue

German: Blaukappentangare

Spanish: Tangara Militar

Taxonomy. Tanagra cyanocephala Statius Müller, 1776, Cayenne; error = Rio de Janeiro, Brazil. Based on molecular-genetic studies this species forms a monophyletic clade with T. fastuosa, T. seledon, T. cyanoventris and T. desmaresti. Three subspecies recognized. Subspecies and Distribution. T. c. cearensis Cory, 1916 – Ceará, in NE Brazil. T. c. corallina (Berlepsch, 1903) – Pernambuco and adjacent N Bahia, in E Brazil. T. c. cyanocephala (Statius Müller, 1776) – SE Brazil (from Espírito Santo S to Rio Grande do Sul), SE Paraguay (Canindeyú) and adjacent NE Argentina (Misiones). Descriptive notes. 13 cm; 16–21·6 g. Unmistakable, brilliantly patterned tanager with green underparts and red nuchal area. Male nominate race has forehead and lores down to chin black, black orbital ring dotted turquoise; upper forehead and ring around eye shining turquoise, crown glistening violet-blue, bright deep red to scarlet band across nape continuing forwards across ear-coverts and cheek, and narrowing onto upper throat; mantle and scapulars deep black, rump and uppertailcoverts bright green; tail feathers dusky, edged and tinged green; lesser upperwing-coverts jetblack, median coverts mostly coppery gold (forming small but conspicuous band), greater coverts black, broadly and extensively edged bright green; basal half of outer flight-feathers and all of inner flight-feathers and tertials broadly edged bright green; small central throat area blackish, becoming violet-blue on lower throat and bright glistening blue across central chest; rest of underparts, including undertail-coverts, bright glistening green; iris dark brown; bill black; legs dark grey to black. Female is similar to male, but much duller (lacking glossiness), has crown dull blue, back greenish with heavy black mottling, nape band duller red, chin black, throat dusky with faint blue tinge. Juvenile resembles adult but much duller. Race cearensis differs from nominate in having crown blue (less violet-tinged), nuchal band less deep red, lower throat cobalt-blue (not violet-blue) and uppertail-coverts glistening turquoiseblue, female like nominate but throat more strongly tinged blue and bordered below by narrow band of brighter blue on chest, belly tinged yellowish-buff; corallina differs from previous in having small patch on throat dark blue, turquoise around eye extends backwards as eyebrow, uppertail-coverts intermediate in colour between those of other races. VOICE. Call a high, sharp “seet!” and softer “sip”, either note sometimes in a series accelerating into a trill; also a high, sharp “tsit-tsit-tsit”. Song possibly a rapid twitter, or the series of notes that quicken to a trill. Habitat. Forest-based species found in canopy, forest borders, light woodland and second growth, and sometimes in urban parks; also occurs in scrub or caatinga in N part of range. To 1000 m or more. Food and Feeding. Fruit supplemented by some insects; reported food items include Phytolacca, melastome berries, Urtica, and myrtle. Occurs in pairs and in lively little parties up to about seven individuals; in forested areas regularly with mixed-species flocks containing others of its genus, but in drier, scrubbier N portions of range (where few such flocks) singles or pairs are usually alone. Searches rapidly, mostly quite high in forest, but will descend low to fruiting shrubs. Breeding. All information from observations of captives. Deep, grassy cup-nest; clutch 3 eggs, whitish with brownish spots, mostly at larger end; incubation entirely by female, period 12–13 days; both adults fed young, which left nest at 15 days and were fed by adults for a further threeand-a-half weeks. Movements. Resident. Status and Conservation. Not globally threatened. Status not well known; common in some places. Range highly fragmented, with four isolated populations, and total numbers now substantially reduced from former levels. Populations once contiguous across S Brazil W to Paraguay and Argentina, but now separate. Still thrives in some numbers in main coastal parts of range, i.e. in Augusto Ruschi (Nova Lombardia) Biological Reserve, in Espírito Santo, and Serra dos Órgãos and Tijuca National Parks, above Rio de Janeiro. In addition, this species is capable of utilizing some second-growth and disturbed habitats. Bibliography. Athanas (2010a), Burns & Naoki (2004), Davis (1945b, 1946), Descourtilz (1852), Diamond & Lovejoy (1985), Dunning (1982), Forbes (1881), Hamilton (1871), Isler & Isler (1999), Lamm (1948), Mitchell (1957), Norgaard-Olesen (1973), Parker & Goerck (1997), de la Peña & Rumboll (1998), Pinto (1954), Ridgely & Tudor (1989, 2009), Sazima et al. (1993), Sedano & Burns (2010), Sick (1985, 1993), Sick & Pabst (1968).

113. Brassy-breasted Tanager Tangara desmaresti French: Calliste de Desmarest German: Orangebrusttangare Other common names: Gould’s Tanager (“T. gouldi”)

Spanish: Tangara de Desmarest

Taxonomy. Tanagra Desmaresti Vieillot, 1819, Rio de Janeiro, Brazil. Based on molecular-genetic studies this species forms a monophyletic clade with T. fastuosa, T. seledon, T. cyanocephala and T. cyanoventris. Described form T. gouldi appears to be a hybrid between present species and T. cyanoventris. Monotypic.

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Distribution. Coastal SE Brazil from SE Minas Gerais, Espírito Santo and Rio de Janeiro S through E São Paulo, E Paraná and E Santa Catarina. Descriptive notes. 13 cm; 19–21·5 g. Colourful, predominantly green tanager with blue on face and large golden patch on throat and chest. Has narrow area around bill and well-defined ocular ring black; otherwise, forehead (rather diffusely demarcated), lores and broad ocular area turquoise, mid-crown to nape, side of neck, mantle and back bright yellowish applegreen, crisply streaked black, rump and uppertail-coverts plain green; tail feathers blackish, broadly edged green; bend of wing and lesser upperwing-coverts brassy golden (often partly hidden), median coverts dusky, broadly edged and tipped brassy golden, greater coverts black, sharply edged and tipped apple-green, flight-feathers black, edged green; side of head green (unstreaked), throat yellow, with rounded black spot in centre of lower throat, below which is a triangular brass-gold area reaching down onto mid-breast; rest of underparts mostly bright green, narrow buff-yellow median line down breast and belly; undertail-coverts mostly yellow, tinged green; iris dark brown; bill black; legs dark horn-grey. Sexes similar. Immature is like adult, but much duller. VOICE. Contact note a high, thin “tseee” or abrupt “tsit!”, singly or repeated rapidly in short series. Also gives rapid chittering trill, and groups may call noisily at times. Habitat. Found mainly in humid pre-montane and montane forest and at forest edge, less often in second growth and clearings with scattered trees. Recorded at 500–2200 m, but mostly at 800– 1800 m. Food and Feeding. Mostly fruits, some arthropods. Reported items include melastome berries, Urtica and myrtle. One stomach contained only vegetable matter. Food items brought to nestlings included spiders (Araneae), seed masses, an orthopteran nymph, caterpillar, planthopper nymphs (Fulgoroidea), katydid nymph (Tettigoniidae) and unidentified winged insect. Markedly social, occurring in pairs, families and monospecific groups of up to about a dozen individuals; frequently found with mixed-species flocks, but moves rapidly and may at times leave slower-moving species behind. Agile, often forages on slender branches within foliage near tip of branch. Of 41 observations of foraging: 16 involved leaf-gleaning; eight were sallies to foliage; seven sallies to air; eight involved scanning and picking at lichens and moss; remainder undocumented. Occasionally forages for arthropods by hopping quickly along limbs, leaning head downwards first on one side of a branch and then on other to inspect surfaces as it progresses outwards; also actively hops and peers in foliage. Breeding. Nest in Jan and another in Feb in Rio de Janeiro (Itatiaia massif). Both sexes bring nest material, but one assumes majority of construction work; nest a deep, thick-walled cup (9 × 8 cm outer diameter) made from dry bamboo leaves (of both native and introduced species), thin twigs, rachises of compound leaves, rootlets, lichen, fungal rhizomorphs, cocoons and spider silk, one nest with moss on outside; 0·5–9 m above ground, one well anchored in multiple-branched fork of tree, two in branched nodes of bamboo, often in man-made clearing. No information on clutch size; eggs whitish, speckled with fine grey spots; male may feed sitting female, incubation period 12–13 days. No other information. Movements. Resident; possibly some movement in Espírito Santo. Status and Conservation. Not globally threatened. Restricted-range species: present in Atlantic Forest Mountains EBA. Locally fairly common. Confined mainly to montane regions where remnant forest patches remain. Occurs in numerous protected areas, among them Serra dos Órgãos National Park, in Rio de Janeiro, Patrimônio Natural Special Reserve, in Minas Gerais, and Itatiaia National Park, on border between those two states. Outside protected areas, this species is regarded as highly vulnerable because of severe forest loss within its limited range. Bibliography. Belcher & Smooker (1937), Bond (1947), Burns & Naoki (2004), Davis (1945b, 1946), Descourtilz (1852), Diamond & Lovejoy (1985), Gagliardi (2007b), Gonzaga & Castiglioni (2006), Hellmayr (1936), Hempel (1949), Holt (1928), Isler & Isler (1999), McCarthy (2006), Meyer de Schauensee (1966, 1970a), Parker & Goerck (1997), Pinto (1944a, 1951), Remsen et al. (2010), Ridgely & Tudor (1989, 2009), Sazima et al. (1993), Sclater (1886), Sedano & Burns (2010), Sick (1985, 1993), Souza (2002), Steinbacher (1938).

114. Gilt-edged Tanager Tangara cyanoventris French: Calliste à ventre bleu German: Blaubrusttangare Other common names: Gould’s Tanager (“T. gouldi”)

Spanish: Tangara Ventriazul

Taxonomy. Tanagra cyanoventris Vieillot, 1819, “Brésil” [Brazil]. Based on molecular-genetic studies this species forms a monophyletic clade with T. fastuosa, T. seledon, T. cyanocephala and T. desmaresti. Described form T. gouldi appears to be a hybrid between present species and T. desmaresti. Monotypic. Distribution. E Brazil from C Bahia (one isolated record from extreme N Bahia), S Minas Gerais and Espírito Santo S to E São Paulo and Rio de Janeiro. Descriptive notes. 13 cm; 16–20 g. Colourful tanager, mostly golden above and blue below, with golden head. Has narrow area surrounding bill and thin orbital ring jet-black; otherwise, entire head, including chin and side of throat, and upperparts golden-yellow, crown, nape and rear side of neck to back streaked black; tail feathers black, edged green; lesser upperwing-coverts mostly bright green (usually concealed), median and greater coverts dusky, edged green, primary coverts and flight-feathers black, edged green, tertials mostly green with dusky inner webs; central throat black, breast and sides turquoise-blue, centre of lower breast and belly yellowish-green, undertail-coverts yellow to yellowish-green or pale yellow-buff; iris dark brown; bill black; legs dark grey. Sexes similar, female tends to be duller. Juvenile undescribed. VOICE. Abrupt sharp call notes include “tsit” and “seet!”, which may accelerate into chittering trill; also a lowerpitched “chip” or “chup”, also sometimes accelerating into short bursts or trills. Habitat. Humid pre-montane forest and forest borders, to lesser extent also second-growth woodland and overgrown clearings. Lowlands to 1200; now mostly above 500 m.

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SPECIES ACCOUNTS Food and Feeding. Diet includes both fruit and some insects. Recorded fruit items include melastome berries and fruit of Urtica. Lives in pairs and in small groups of up to about eight individuals, often found with mixed-species flocks containing others of genus; possibly not so often in large monospecific groups as is T. desmaresti. Forages from middle heights up to canopy in forest, but descends lower along forest edges to exploit fruiting shrubs. Searches for insects mostly by hopping among small slender outer twigs and peering in terminal foliage; occasionally also checks curled green leaves. Behaviour much like that of forest-based congeners of SE Brazil. Breeding. A nest found in Nov in Rio de Janeiro was 12 m up in fork of tree; nest possibly lined with strips of bamboo. Offspring of a previous brood have been seen to act as helpers at nest. No other information.

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PLATE 11 Movements. Resident. Status and Conservation. Not globally threatened. Locally common. Range now much fragmented, and numbers substantially reduced. Survives in remaining forested areas in SE coastal range, and (as with many other SE Brazil endemics) bulk of population may now be confined largely to protected areas. Considered still common at several reserves, including Augusto Ruschi (Nova Lombardia) Biological Reserve, in EspĂrito Santo, and Itatiaia National Park, in Rio de Janeiro. Bibliography. Dunning (1982), Erickson & Mumford (1976), Hellmayr (1936), Holt (1928), Isler & Isler (1999), McCarthy (2006), Meyer de Schauensee (1966, 1970a), Mitchell (1957), Parker & Goerck (1997), Pinto (1951), Remsen et al. (2010), Ridgely & Tudor (1989, 2009), Sclater (1886), Sedano & Burns (2010), Sick (1985, 1993), Skutch (1961), Souza (2002).

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Class AVES Order PASSERIFORMES Suborder OSCINES

Family EMBERIZIDAE (BUNTINGS AND NEW WORLD SPARROWS) • Tiny to medium-sized passerines with conical bill, many simply patterned in brown, black, grey, yellow and white, often streaked brown or grey, some more colourful and/or strikingly marked. • 8·5–24 cm.

• Holarctic, Neotropical, Afrotropical and Oriental Regions. • Open and semi-open habitats, especially grassland; also woodland and forest edge. • 76 genera, 326 species, 887 taxa. • 36 species threatened; at least 3 subspecies extinct since 1600.

Systematics The buntings and the New World sparrows form a large assemblage of passerines with a geographical range that covers most of the world. They are absent only from the Australasian Region and Oceania. As currently constituted, the family Emberizidae contains more than 300 species, 326 in the present treatment, but it seems highly likely, on the basis of recent molecular-genetic research, that several genera do not really belong in this family. Although the family has a wide distribution, the majority of its members, more than 80%, are found in the New World, and it is in this part of the world that most of the generic diversification has occurred. They are closely related to a larger group, commonly referred to collectively as the “New World nine-primaried oscine passerines”, a not entirely accurate designation because some species are not exclusively New World taxa, and some other oscine groups that have only nine primary wing feathers are not placed in this group. The New World nine-primaried oscines are generally divided into as many as seven families, and most of the species seem to fall within one or other of these. The families include the New World warblers (Parulidae), with 116 species; the tanagers (Thraupidae), containing 283 species in the present treatment; the cardinals (Cardinalidae) with 42 species; the New World blackbirds (Icteridae), containing 111 species; and the present family, Emberizidae, with more than 300 species. Some authors recognize Coerebidae, with the Bananaquit (Coereba flaveola) usually as its the sole member, and recently some separate the longspurs and snow buntings as the family Calcariidae, with a total of six species. In the present arrangement, the Bananaquit is included within the tanager family, while the longspurs and snow buntings are incorporated within Emberizidae. With the exception of Emberizidae (including Calcariidae), all of these families normally occur only in the New World. Collectively, they contain nearly 880 species, some 8% of the world total. The Olive Warbler (Peucedramus taeniatus), which has generally been accorded its own monotypic family, Peucedramidae, has sometimes been placed with the above families because it resembles a New World warbler, but it appears not to be closely allied to them. Indeed, molecular analysis carried out by K. A. Jønsson and J. Fjeldså suggested that Peucedramus is allied to the Eurasian accentors (Prunellidae).

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Most authors now recognize five or six basic groups of taxa, the Bananaquit being probably best placed with the tanagers, rather than in a monotypic family. They differ, however, in their hierarchical classifications, some recognizing most or all of the aforementioned families as subfamilies within a greatly expanded Emberizidae. For example, E. C. Dickinson, in the revised and enlarged third edition of The Howard and Moore Complete Checklist of the Birds of the World, published in 2003, treats the groups as separate families, as also does the AOU Check-list of North American birds. C. E. Hellmayr, in the first half of the twentieth The three longspur species in the genus Calcarius are not closely related to other members of Emberizidae. Together with McCown’s Longspur (Rhynchophanes mccownii) and the Snow Bunting (Plectrophenax nivalis) and McKay’s Bunting (P. hyperboreus), they are sometimes placed in a new family, Calcariidae. Smith’s Longspur (Calcarius pictus) and the Chestnut-collared Longspur are exclusively North American in their distribution, but the Lapland Longspur (C. lapponicus) breeds throughout the arctic and subarctic regions of North America, Europe and Asia. Longspurs are named after their elongated hind claw. [Calcarius ornatus, Saskatchewan, Canada. Photo: Glenn Bartley]

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The Snow Bunting is a large emberizid with a large conical bill and relatively short, notched tail. The wings are pointed and relatively long, which is typical of emberizids that migrate long distances or live in windy environments. The three Calcarius longspurs include two species with long, pointed wings and one with shorter and more rounded wings. Snow Buntings are sexually dimorphic both in their breeding plumage and in size, the female being smaller. Geographical variation in Snow Buntings is not well understood and is complicated by a high degree of individual variation. [Plectrophenax nivalis nivalis, Helsinki, Finland. Photo: Markus Varesvuo]

century, treated the cardinals, Darwin’s finches of the Galapagos Islands, the cardueline finches, and the New World sparrows and buntings as subfamilies in the family Fringillidae, whereas C. G. Sibley and B. L. Monroe, in 1990, considered them to be distinct tribes within the subfamily Emberizinae. Shortly thereafter, S. Cramp and C. M. Perrins, in the Handbook of the Birds of Europe, the Middle East and North Africa, listed Parulidae, Thraupidae and Icteridae as three separate families, and split the Emberizidae into the subfamilies Emberizinae and Cardinalinae. Although these basic groups are generally recognized by all taxonomists, there are many species that appear to be borderline cases. As a result, a relatively large number are considered to be incertae sedis, of uncertain position, this no doubt reflecting the close similarity among the species of nine-primaried oscines. As more information is gathered and more knowledge gained, especially through molecular biology, it is certain that substantial changes in the higher-level classification will continue to be made. Over the past century, much information has been gathered by ornithologists with regard to the affinities of these nineprimaried oscines. Although explicit criteria for classification usually were not stated, traditional taxonomies were doubtless based on external morphology, especially bill configuration: buntings and New World sparrows generally have a conical bill that can be used for cracking and milling seeds. Characters associated with feeding, however, seem to be evolutionarily malleable, and subject to convergence. We have learnt that many species with bills that are well suited for seed-eating, or, for that matter, for nectarivory or fruit-eating, have apparently evolved independently in groups that are otherwise not particularly closely related; bill morphology alone, therefore, does not necessarily predict similarities based on other features. In other words, many bird species with a conical bill like that of a New World sparrow are not particularly closely related to the Emberizidae. Some studies have concentrated on various anatomical and physiological features. In particular, W. J. Beecher studied comparative jaw musculature; D. Mainardi, C. G. Sibley and W. Stallcup examined pelvic musculature and serology; J. Robins, G. Schnell, J. D. Rising, J. D. Webster and J. R. Webster undertook multivariate analyses of skeletal measurements; H. B. Tordoff investigated cranial and palatal characters; and R. Raikow studied appendicular myology. All of this research has generally supported the recognition of the Emberizidae as a natural group which, if not monophyletic,

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is at least paraphyletic, but it has yielded few characteristics that have supported a constant classification. Ongoing molecular analyses are contributing significantly to an understanding of the relationships among these nine-primaried oscine species. In 2006, for example, Jønsson and Fjeldså compared results from 99 different molecular analyses of sparrowlike birds and provided a useful summary. More recently, analyses of mitochondrial DNA by J. M. DaCosta and colleagues, added to earlier work, clarified the relationships of the species within the genera Aimophila and Pipilo, supporting the contention that these genera are, as delimited in earlier taxonomies, polyphyletic. Again, these studies corroborate the conclusions of the many ornithologists who had previously based their analyses on morphology, breeding biology, ecology and behaviour. Modern molecular-genetic research has strongly indicated that several genera placed in the present family Emberizidae almost certainly do not belong there. The monotypic genus Euneornis, formerly placed in Coerebidae, a family no longer recognized, and currently included in Emberizidae, may well be closest to the tanagers in the family Thraupidae. Similarly, several other Caribbean genera, namely Loxipasser, Melopyrrha, Loxigilla and Melanospiza, are probably better included within Thraupidae, as also are the Galapagos genera Certhidea, Platyspiza, Camarhynchus and Geospiza. In fact, these modern molecular-genetic studies suggest that 19 other genera currently in Emberizidae are more closely related to members of the tanager family. These are the Tiaris grassquits, the high-Andean and Patagonian Phrygilus and Diuca, Haplospiza, the Poospiza mountain-finches of the high Andes, Sicalis yellow-finches, Embernagra pampa-finches, the monotypic grassquit genus Volatinia, the Sporophila seedeaters and Oryzoborus seed-finches, the Catamenia seedeaters of the high Andes, the Central American Acanthidops, the Argentine Saltatricula, the Coryphospingus and Rhodospingus finches, the Paroaria cardinals, and the three monotypic genera Porphyrospiza, Charitospiza and Gubernatrix. The Snow Bunting (Plectrophenax nivalis) and the Lapland Longspur (Calcarius lapponicus) are Holarctic in distribution, but probably of New World origin. In North America there are two species of Plectrophenax, one of which, McKay’s Bunting (Plectrophenax hyperboreus), breeds only on a few islands in the Bering Sea, and is perhaps better treated as a well-marked race of the Snow Bunting. Of the four species of longspur, only the Lapland Longspur has an Holarctic distribution, the other

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All except four of the 35 Eurasian emberizid species are, at present, placed in the genus Emberiza. The Yellowthroated Bunting’s permanently erect crest is unusual; most emberizids with a crest raise this only when excited or displaying. The House Bunting is more typical of Emberiza (and, with its rather plain plumage, easily confused with three other Emberiza species in its range), though smaller and slighter than many of its congeners. [Left: Emberiza elegans elegantula, Sichuan, China. Photo: Zhou Huaming. Right: Emberiza sahari, Oued Massa, Morocco. Photo: Mathias Schäf]

three being exclusively North American. The Lapland Longspur, until fairly recently considered an Emberiza in Europe, has a circumpolar distribution, breeding throughout arctic and subarctic regions; Smith’s Longspur (Calcarius pictus) breeds in northern Canada and central Alaska, and McCown’s Longspur (Rhynchophanes mccownii) and the Chestnut-collared Longspur (Calcarius ornatus) breed in the northern Great Plains. Molecular studies undertaken by R. Carson and G. Spicer, T. Yuri and D. Mindell, Jønsson and Fjeldså, J. Klicka and co-workers, and P. Alström and colleagues suggest that these six species are not closely related to other members of Emberizidae, and may, in fact, be basal to many of the other New World nine-primaried oscines. Some taxonomists nowadays place them in a new family, Calcariidae. While such treatment may prove to be preferable, the present treatment includes these “calcariid” genera in Emberizidae, as has been the traditional arrangement. Indeed, in terms of their biology and in various other aspects, they are very similar to the emberizids. Of the 326 species in the Emberizidae as currently constituted, 35 are found in Eurasia. All except four of these are, at present, placed in the genus Emberiza. The exceptions are the Lapland Longspur and Snow Bunting, both found across the Holarctic; the Crested Bunting (Melophus lathami), resident in southern Asia from the Indian Subcontinent eastwards to southeast China; and the Slaty Bunting (Latoucheornis siemsseni), restricted to central and eastern China. The Corn Bunting (Emberiza calandra) of Europe and western Asia was for a while generally placed in the monotypic genus Miliaria, but in recent years it has been returned to Emberiza, in which it had previously been included. All nine African members of the family are included in the genus Emberiza. In the South Atlantic, the Gough Finch (Rowettia goughensis) is confined to the island of that name in the Tristan da Cunha group, and the three species of Nesospiza are likewise found only in that group of islands, the Inaccessible Finch (Nesospiza acunhae), the Nightingale Finch (Nesospiza questi) and Wilkins’s Finch (Nesospiza wilkinsi) being restricted to, respectively, Inaccessible Island, Nightingale Island and some of its offshore islets, and Nightingale Island alone; the three are sometimes treated as conspecific. All of these Tristan species are probably derived from South American emberizids. The Gough Finch, for example, closely resembles the Canary-winged

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Finch (Melanodera melanodera) of the Falkland Islands, and there are strong prevailing westerly winds at that latitude. The genus Emberiza is not represented in the Americas, except in the form of rare migrants and vagrants from the north-east Palearctic Region. No species of Emberizidae is native to the Australasian Region. The sparrows inhabiting the northern temperate region of America have often been divided into two general groups, the grassland sparrows and the brushland sparrows. This separation, while based on ecological and behavioural characteristics, seems also, in general, to separate groups of closely related species, although several genera are not easily placed in either category. The grassland clade has been thought to include Passerculus, Ammodramus and, perhaps, Xenospiza. Of these, Passerculus, the savannah-sparrow complex, has often been subsumed in Ammodramus, but molecular data do not support this treatment. Similarly, the monotypic Xenospiza, containing the Sierra Madre Sparrow (Xenospiza baileyi), has been placed in Ammodramus, but this has been questioned for several reasons. F. A. Pitelka, for example, argued that the similarities of Xenospiza to Ammodramus were superficial, and that, in general size and proportions, the Sierra Madre Sparrow was more like Lincoln’s Sparrow (Melospiza lincolnii). Klicka and G. M. Spellman’s molecular data separate Ammodramus into two groups, a grassland group and a marshland group. The former included the savannahsparrow complex, currently in Passerculus, and Baird’s Sparrow (Ammodramus bairdii), the Grasshopper Sparrow (Ammodramus savannarum), the Grassland Sparrow (Ammodramus humeralis) and the Yellow-browed Sparrow (Ammodramus aurifrons). The marshland group incorporated the Seaside (Ammodramus maritimus), Nelson’s (Ammodramus nelsoni), Saltmarsh (Ammodramus caudacutus), Le Conte’s (Ammodramus leconteii) and Henslow’s Sparrows (Ammodramus henslowii), and perhaps also the Sierra Madre Sparrow. Robins and Schnell’s multivariate analysis of 48 skeletal characters of selected species of grassland sparrow more or less supports this separation, placing the Grassland, Grasshopper and Baird’s Sparrows and the savannahsparrow complex in one group, and Le Conte’s, Saltmarsh and Seaside Sparrows in another; Xenospiza is in the former group, but does not cluster closely with it, and Henslow’s Sparrow is with the latter, but again is not closely clustered with it. Within the grassland assemblage, the Grassland Sparrow and the Yellow-

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The “Fox Sparrow” was formerly treated as a single, highly variable species. The 18 recognized races were divided into four groups according to plumage, bill morphology and breeding distribution: red (iliaca), sooty (unalaschcensis), thickbilled (megarhyncha) and slate-colored (schistacea). Mitochondrial DNA analysis provided support for the four groups to be recognized as species, which is the treatment followed here. The race zaboria of the Red Fox-sparrow (shown here) is more faintly streaked than the nominate, but plumage varies gradually over distance, and many individuals cannot safely be assigned to either race. [Passerella iliaca zaboria, British Columbia, Canada. Photo: Glenn Bartley]

browed Sparrow occur only in South America, and have sometimes been put into a separate genus, Myospiza. From recent molecular studies, including that by Klicka and Spellman, and others, it appears that Henslow’s Sparrow and Baird’s Sparrow are sister-species, and Le Conte’s Sparrow, the Saltmarsh Sparrow, Nelson’s Sparrow and the Seaside Sparrow form a monophyletic clade. Traditionally, the savannah-sparrow complex, the Grassland Sparrow and the Yellow-browed Sparrow have been thought to be closely related to Henslow’s and

Baird’s Sparrows, but the molecular data available do not support this. Klicka and Spellman, however, found that the Sierra Madre Sparrow clusters with the three species of Melospiza, namely the Swamp Sparrow (Melospiza georgiana), Lincoln’s Sparrow and the Song Sparrow (Melospiza melodia), which have traditionally been placed with the brushland sparrows, and these, in turn, cluster with Henslow’s Sparrow, Baird’s Sparrow and the savannah-sparrow complex. Their analyses also support the placement of the Vesper Sparrow (Pooecetes gramineus) and Bell’s Sparrow

Recent molecular-genetic studies suggest that the genus Junco spread quickly north from Central America in the postglacial period. The farther south in Central America, the older is the lineage. The southernmost, and the largest of the three species, is the monotypic Volcano Junco. Its closest relative is the southern race alticola of the Yellow-eyed Junco (Junco phaeonotus). Northern populations of Yellow-eyed Junco are, in turn, very close genetically and morphologically to southern races of the Dark-eyed Junco (J. hyemalis). The 16 races of the Dark-eyed Junco vary in size, plumage and bill colour and form six groups, which are sometimes treated as six separate species. [Junco vulcani, Irazú Volcano National Park, Costa Rica. Photo: Boris Nikolov]

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(Artemisiospiza belli), but not the Black-throated Sparrow (Amphispiza bilineata), and the monotypic Striped Sparrow (Oriturus superciliosus) appears to belong in this group. Recent work shows that the Five-striped Sparrow (Amphispiza quinquestriata), formerly included in the genus Aimophila, is best placed in Amphispiza, close to the Black-throated Sparrow. What was formerly a large genus, Aimophila, has recently been split into several smaller genera. Its greatest species rich-

ness was in Mexico and Central America, a few species reaching the southern USA, and with two found only in South America. Research by L. Wolf, however, indicated that Aimophila was not monophyletic, and that its species should be separated into three or four groups that probably have separate evolutionary histories and are not particularly closely related to each other. This has recently been supported by the genetic work of DaCosta and coworkers, and their results have been widely accepted by the ornithological community. Two groups of species are now placed in the genus Peucaea: the Cinnamon-tailed (Peucaea sumichrasti), Rufous-winged (Peucaea carpalis), Stripe-headed (Peucaea ruficauda), Black-chested (Peucaea humeralis) and Bridled Sparrows (Peucaea mystacalis); and Cassin’s (Peucaea cassinii), Bachman’s (Peucaea aestivalis) and Botteri’s Sparrows (Peucaea botterii). These all occur in grassland, often xeric, and among xeric shrubs, and in pine (Pinus) savanna in Mexico and the southern USA. The first group, the Cinnamon-tailed, Rufouswinged, Stripe-headed, Black-chested and Bridled Sparrows, probably radiated in the lowland thorn-forests of Mexico and the Pacific lowlands of Central America. The third group contains the Rufous-crowned (Aimophila ruficeps), Rusty (Aimophila rufescens) and Oaxaca Sparrows (Aimophila notosticta), which appear to have radiated in the pine–oak (Pinus–Quercus) woodlands of Mexico and Central America; these remain in the genus Aimophila, the type species of which is Aimophila rufescens. As mentioned above, the Five-striped Sparrow, often placed in Aimophila, is now included with the Black-throated Sparrow in Amphispiza, and this genus appears to be closely allied to Chondestes and Calamospiza. Recent genetic data published by DaCosta and colleagues showed that the relationships among some of the North American genera were not properly reflected in the linear sequence of earlier lists. For example, the towhee genus Pipilo was diphyletic. Four species, the Collared (Pipilo ocai), Green-tailed (Pipilo chlorurus), Spotted (Pipilo maculatus) and Eastern Towhees (Pipilo erythrophthalmus), should follow Atlapetes in the sequence, with the three Aimophila species, the Rufous-crowned, Rusty and Oaxaca Sparrows, coming next. The four species of “brown towhee”, namely the Canyon (Pyrgisoma fuscum), Whitethroated (Pyrgisoma albicolle), California (Pyrgisoma crissale) and Abert’s Towhees (Pyrgisoma aberti), are now placed in the

The sparrows inhabiting the northern temperate region of America have often been divided into two general groups, the grassland sparrows, to which Ammodramus belongs, and the brushland sparrows. Molecular data have been used to separate Ammodramus into two further groups, a grassland group and a marshland group, the latter containing the Seaside Sparrow along with Nelson’s (Ammodramus nelsoni), Saltmarsh (A. caudacutus), Le Conte’s (A. leconteii) and Henslow’s Sparrows (A. henslowii). Seven extant races of Seaside Sparrow are recognized. Two races, including the very distinctive “Dusky Seaside Sparrow” (nigrescens), are extinct. [Ammodramus maritimus fisheri, Brazoria, Texas, USA. Photo: Brian E. Small]

In keeping with their largely terrestrial lifestyles, the New World sparrows are generally brownish or rusty in colour. Many, including the Lark Sparrow, have a boldly marked head pattern. The tail pattern of the Lark Sparrow is also very distinctive, the outermost pair of rectrices being edged with white, and the other outer rectrices boldly white-tipped. The sexes are similar in coloration, although the female is somewhat smaller. The Lark Sparrow occupies the monotypic genus Chondestes. It appears closest to the Lark Bunting (Calamospiza melanocorys) and the Five-striped (Amphispiza quinquestriata) and Black-throated Sparrows (A. bilineata). [Chondestes grammacus strigatus, Alberta, Canada. Photo: Glenn Bartley]

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Family EMBERIZIDAE (BUNTINGS AND NEW WORLD SPARROWS) Until recently, the Sage Sparrow and Bell’s Sparrow (Artemisiospiza belli) were placed in the “brushland” sparrow genus Amphispiza, but recent molecular work indicates that the two are, instead, members of a well-resolved clade of “grassland” sparrows. They are, however, found in sagebrush, chaparral and other scrub habitats. Bell’s and Sage Sparrows were formerly considered conspecific, the Sage Sparrow differing mainly in its larger size, paler coloration and more obvious streaking above. The race canescens of Bell’s Sparrow, which is very similar to the Sage Sparrow and possibly hybridizes with it where the ranges meet, may be a separate species. [Artemisiospiza nevadensis nevadensis, Mono County, California, USA. Photo: Brian E. Small]

The Black-chested Sparrow is a restrictedrange species from Mexico. Two of its closest relatives are also Mexican endemics with restricted ranges. The Peucaea species were formerly included in Aimophila, a large genus of mostly Mexican and Central American sparrows of arid tropical scrub-forest and thornforest habitats. The Aimophila species were morphologically and vocally diverse, and held by many to constitute an “unnatural assemblage”. Genetic data subsequently indicated that Aimophila included four distinct lineages. The resurrected genus Peucaea includes eight species. [Peucaea humeralis astica, Colima, Mexico. Photo: Pete Morris]

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genus Pyrgisoma along with the Rusty-crowned Ground-sparrow (Pyrgisoma kieneri), which was formerly placed with the two other ground-sparrows in the genus Melozone. The summary analyses of Jønsson and Fjeldså suggest that the Chlorospingus bush-tanagers, conventionally classified with the tanagers in Thraupidae, belong in this group. The grassquits in the genera Volatinia and Tiaris, the seedeaters in Sporophila, the seed-finches in Oryzoborus, the grass-finches in Emberizoides, and their allies form a widespread and diverse group of small, seed-eating Neotropical nineprimaried oscines. Although they have often been included within Emberizidae, Sibley and Monroe placed them with the tanagers in Thraupidae, and this treatment is supported by the moleculargenetic studies of Jønsson and Fjeldså. The Caribbean bullfinches (Loxigilla, Melopyrrha), the Yellow-shouldered Grassquit (Loxipasser anoxanthus), the Orangequit (Euneornis campestris) and the St Lucia Black Finch (Melanospiza richardsoni) are part of this group, as probably are the Galapagos finches in the genera Geospiza and Camarhynchus. These molecular data also indicate that the South American cardinals (Paroaria), flowerpiercers (Diglossa), sierra-finches (Phrygilus) and warbling-finches (Poospiza) should all be placed in the tanager family. Unsurprisingly, a number of taxonomic problems at the species level remain to be resolved. For example, in the genus Sporophila, the second largest in the family after Emberiza, some authors recognize a species Sporophila melanops, given the English vernacular name of the “Hooded Seedeater”. Indeed, this is even listed “officially” as a Critically Endangered emberizid. It is known only from a single specimen, an adult male taken in October 1823 on the western border of Goiás, in Brazil; the precise locality was a lake 15 km north of Registro do Araguaia, on the east bank of the Araguaia River. This seedeater has not been found again, even with the advantages of modern technology and methods. Searches carried out along the floodplain of the Araguaia, most recently in December 2008 and January 2009 and, again, in July 2010, failed to produce any sightings. Although the Hooded Seedeater is included in a number of checklists and other authoritative works, it is considered likely that it was, in fact, a variant of the Yellow-bellied Seedeater (Sporophila nigricollis) or perhaps a hybrid between it and another member of this genus. Another seedeater of uncertain standing is the “Tumaco Seedeater”, described as Sporophila insulata and known only from the island of Tumaco, a short distance off the coast of the south-west Colombian province of Nariño. It has a rufous rump and rufous underparts, but is otherwise more or less identical to the Chestnut-throated Seedeater (Sporophila telasco), which is resident on the Pacific coast and coastal slope from south-west Colombia southwards through the coastal lowlands to the desert valleys of northernmost Chile. The Tumaco population, which may well be extinct, is thought to be a subspecies or a variant of the Chestnut-throated Seedeater or, perhaps less likely, a hybrid between the latter and the Ruddy-breasted Seedeater (Sporophila minuta). A further example from this problematic genus involves the form described as Sporophila zelichi, known as “Narosky’s Seedeater” or the “Entre Ríos Seedeater”, the latter name denoting the region of north-east Argentina from where the bird was initially described, in 1977. Narosky’s Seedeater is still poorly known. It differs from the Marsh Seedeater (Sporophila palustris) only in its possession of a broad white nuchal collar, and is indistinguishable from the latter in its vocalizations and habitat choice; the two can even be found at the same localities. The form “zelichi” is considered to be a white-necked morph of the Marsh Seedeater or just possibly a hybrid between it and the Chestnut Seedeater (Sporophila cinnamomea). Hybridization is probably not uncommon among the emberizids. Indeed, the Yellowhammer (Emberiza citrinella) and the Pine Bunting (Emberiza leucocephalos) regularly interbreed in a fairly extensive area of western and central Siberia, and there are many examples of occasional hybridization involving other members of the family. Among Darwin’s finches in the Galapagos Islands, the Medium Ground-finch (Geospiza fortis) interbreeds rarely with the Small Ground-finch (Geospiza fuliginosa), and the resultant hybrids tend to have low fitness. On the island of

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Daphne Mayor, however, following the 1982–1983 El Niño event and the accompanying huge rainfall, these smaller-billed hybrids had elevated fitness as a result of the subsequent superabundance of small seeds on the island (see also Morphological Aspects). Similarly, hybridization between the Medium Ground-finch and the Common Cactus-finch (Geospiza scandens) is rare and hybrids have low fitness, but the latter’s fitness was again greater after the El Niño event. It is thought that large-scale hybridiza-

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HANDBOOK OF THE BIRDS OF THE WORLD lebruni), but it was then reidentified as an isolated form of the Greater Yellow-finch (Sicalis auriventris). However, quite apart from its notable geographical separation from the otherwise known ranges of both of these species, the big problem is that its nesting behaviour is quite different from both of theirs. Its cupnest, built from dry grasses and with a finer lining, is placed approximately 36 cm up in tussock paspalum grass (Paspalum quadrifarium); in this it lays four whitish eggs, with darker markings concentrated at the wide end, which it incubates for twelve days, and the young take at least ten days to fledge. The nest-site is very different from those of both of the species in which it has been tentatively placed, and clutch size and egg coloration are also different from those of the Greater Yellow-finch. It could be that an as-yet-undescribed species is involved.

Morphological Aspects

tion between the Medium Ground-finch and the Large Groundfinch (Geospiza magnirostris) has occurred on Santa Cruz, where the former species is noticeably larger-billed than are its populations on other islands in the archipelago. Several intriguing taxonomic problems require full investigation. As just one example, in Argentina, a form of yellow-finch breeding in the Sierra de la Ventana, in Buenos Aires, was originally identified as the Patagonian Yellow-finch (Sicalis

Buntings and New World sparrows are tiny to medium-sized songbirds, ranging in mass from about 6 g to almost 70 g, and in length from about 8·5 cm to 25 cm. As a group, the smallest emberizids are the grassquits and seedeaters, 6–15·5 g in weight and 8·5–12 cm in length. The Yellow-bellied Seedeater, for instance, has a length of 8·5–10·3 cm and weighs 8·5–11·2 g, the Ruddy-breasted Seedeater measures 8·6–9·6 cm and weighs 7– 9 g, and the corresponding figures for the Blue-black Grassquit (Volatinia jacarina) are 8·7–10·9 cm and 8–12 g. Sparrows in the genus Spizella are among the smallest of the New World sparrows, generally being 10–15 g in weight and 11–14 cm in length. The largest of the sparrows are the towhees: the California Towhee is 21–24 cm in length and weighs about 33–67 g. Weight data are, of course, of limited value in expressing size, because many individuals, prior to migration, deposit large quantities of fat, sometimes even doubling their usual body mass. In addition, the body mass of a females during the laying period varies considerably and on a daily basis. Many species exhibit geographical variation in size. Song Sparrows, for example, provide a striking example of this. Wing lengths of males of the subspecies maxima and sanaka, from the Aleutian Islands, in Alaska, range from 82 mm to 87 mm, females being slightly smaller, 78–85 mm, whereas, at the other

The Neotropical sparrow genus Arremon includes more than 20 mediumsized, robustly built, predominantly greenish and relatively colourful species. The Goldenwinged Sparrow is a stocky and pot-bellied, long-billed and shorttailed sparrow, one of a group of five similar species that may form a superspecies. Two former genera, Lysurus and Buarremon, were merged with Arremon in 2008. The eight taxa forming the “Whitebrowed Brush-finch complex” (formerly Buarremon torquatus) were subsequently proposed as representing eight distinct species, a treatment followed here. [Arremon schlegeli schlegeli, Santa Marta, Colombia. Photo: Pete Morris]

The ground-foraging Large-footed Finch uses its large feet and powerful legs to scratch in soil and leaf litter. The simultaneous use of both feet, bilateral or double scratching, is widespread among the Emberizidae, but especially well developed in this species. The Large-footed Finch is the only species in the genus Pezopetes, but is sometimes included in the brush-finch genus Atlapetes. It typically lays only one egg, which is unusual for the family in general and for mainland emberizids in particular. [Pezopetes capitalis, San Gerardo, Costa Rica. Photo: Theodoulos Poullis]

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Family EMBERIZIDAE (BUNTINGS AND NEW WORLD SPARROWS)

The brush-finches of the genus Atlapetes have their greatest diversity in the Andes. Forms with different colours may replace each other sharply at different altitudes on the same slopes, or in similar habitats on different slopes. In older classifications, grey species such as the Slaty Brush-finch and yellow-and-green species such as the Bolivian Brush-finch were placed in different groups, and similarly coloured populations replacing each other in corresponding ecological zones on different slopes were ranked as subspecies of the same species. Mitochondrial DNA sequence data indicate that traditional groupings based on plumage colours did not follow evolutionary trajectories in the genus, and also that many forms currently ranked as subspecies are genetically more divergent than sympatric species. The restrictedrange Santa Marta Brush-finch is the only Atlapetes species in the Santa Marta Mountains of Colombia, but the Tumbesian endemic White-winged Brushfinch is found in mixed flocks with other brush-finches. [Top left: Atlapetes schistaceus schistaceus, Chicaque, Colombia. Photo: Jussi Vakkala. Top right: Atlapetes rufinucha rufinucha, Carrasco National Park, Cochabamba, Bolivia. Photo: Daniel Alarc贸n. Bottom left: Atlapetes melanocephalus, El Dorado Bird Reserve, Colombia. Photo: Ketil Knudsen. Bottom right: Atlapetes albiceps, Chaparri Ecological Reserve, Tumbes, Peru. Photo: Heinz Plenge]

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