Condor, 82:379-391 0 The Cooper Ornithological Society 1980

THE OF FINCA MERENBERG, HUILA DEPARTMENT,

ROBERT S. RIDGELY AND STEVEN J. C. GAULIN

ABSTRACT.-An annotated list of the birds found in a privately owned re- serve in the subtropical zone on the eastern slope of Colombias’ Central is presented. The list is based on our own observations carried out in 1975-1976, and is amplified by discussion of hitherto unreported collections of birds taken nearby by M. A. Carriker, Jr., in the 1950s. Included are dis- tributional, behavioral, and ecological notes on a number of poorly known . Important range extensions are noted for four species: Hapalopsit- taca amaxonina, Anthocephalafloriceps, pucheranii, and Atlapetes Javiceps.

Colombia possesses one of the worlds’ most specimens at about 2,500 m in Moscopan, diverse avifaunas. This is in large part due but almost none of this material has been to the tremendous habitat diversity con- reported on in the literature. Major holdings tained within its borders. Detailed ornitho- of these collections now reside in several logical studies of particular areas are, how- U. S. museums (National Museum of Natu- ever, rare, and major recent works on ral History, Field Museum of Natural His- Colombian birds (Meyer de Schauensee tory, Yale Universitys’ Peabody Museum of 1948-1952, 1964) are not organized so as to Zoology, Carnegie Museum, and Louisiana present a picture of local avian communi- State University Museum of Zoology) and ties. Chapman (1917) did discuss overall are discussed below in relation to our own distribution, but this was his sole focus; findings. for lack of information he could say little about ecology and behavior, and much of DESCRIPTION OF THE STUDY AREA his work is now nomenclaturally or taxo- Our principal study site was Finca Meren- nomically obsolete. Our study focuses on berg, a privately owned reserve and farm the birds of a small area in Moscopan, De- situated 50 km west of La Plata on the Po- partment of Huila, Colombia. We report the payan-Neiva highway (2”14N;’ 76”8W).’ species present, their abundance, micro- The reserve encompasses 270 ha on the habitat preferences and migratory status, eastern slope of the Central Cordillera at an and present new data on their ecology, be- average elevation of 2,300 m; about 200 m havior and distribution. separate the lowest and highest points with- Moscopan was first referred to in the or- in the site. It falls within Holdridges’ (1967) nithological literature by Meyer de “lower montane wet ” zone. Schauensee (1948) who characterized it as About half of the reserve is forested, most- “2,000-2,900 m Huila. The name of a ly the steeper slopes, which contain diverse stream and of the general region of the up- tree species. The most common genera are per Rio La Plata Valley, 32 km west of the Landerbergia, Guarea, Cecropia, Morus, town of La Plata, 2”20N.”’ The Rio La Plata Ficus, Hieronyma, Billia, Turpinia, Sap- is a tributary of the drain- ium, Nectandra and Quercus. Some indi- ing a portion of the eastern slope of the Cen- viduals of Ficus sp. and Quercus humbold- tral Cordillera of the Andes in the geograph- tii attain a height of 40 m and a diameter at ically complex region near the Magdalenas’ breast height of 1.5 m. In the understory, source. Previous ornithological studies in the tree fern Dryopteris and the arbores- this region are virtually nonexistent: Leh- cent shrubs Miconia and Palicourea are mann (1957) did publish notes on a few common. Where sunlight reaches the forest species he collected in the area in the floor, the undergrowth is very dense, the 1950s. Chapmans’ (1917) nearest locality bamboo Chusquea being a major element. was La Candela, situated in a different wa- Most trees are covered with vines, orchids, tershed to the south at 2,000 m. During the bromeliads, and other epiphytic . The 195Os, M. A. Carriker, Jr., collected over 500 remaining, more level, half of the reserve 13791 380 ROBERT S. RIDGELY AND STEVEN J. C. GAULIN

TABLE 1. Meteorological data from Finca Meren- foods being consumed and to record the foraging style. berg. In the case of large mixed flocks, which were frequent, complete data could not always he recorded for every Temperature (“C) species. The analysis consisted of compiling both the forest and edge records so as to assemble a picture of A\WMge Average daily daily Rainfall the relative abundance of every species and to char- Month maximum mmimum (mm) acterize each in terms of its microhabitat preferences, behavior, diet, and grouping tendencies. Certain small January 17.5 11.1 49 understory species were particularly hard to identify February 17.0 11.1 159 and thus, to complement our observational records, we March 17.3 11.6 270 operated mist-nets in the forest and at the edge for April 17.7 11.5 264 three days in mid-May. All hut 2 of the 89 individuals May 18.3 11.3 239 trapped were subsequently released. At the three suh- June 17.3 11.1 240 sidiary sites we used the same techniques, hut mist- July 15.6 10.2 182 netted only at Cueva de 10s Guicharos National Park. August 122 In the species accounts, five categories of abundance September 93 have been used: &t&ant-recorded daily, often in October 153 large numbers; common-recorded almost daily, hut November 94 not usually in large numbers; uncommon-recorded December 76 on 20-60% of days afield, rarely if ever in large num- Total 1,942 hers; rare-recorded only occasionally; and oery rare-recorded less than five times, some species being vagrants while others were presumably resident hut exceptionally rare or inconspicuous. Species consists of pastures of sown grasses, but low marked with an asterisk were collected by Carriker in intensity grazing has allowed partial recol- Moscop6n (see below). and nomenclature, onization by native plants in some areas. with a very few exceptions, follow Meyer de Schauen- The pastures also contain isolated trees see (1966, 1970). which were not cut during the original SPECIES ACCOUNTS clearing. Corugyps utrutus. Black Vulture. Common over clear- Meteorological data were not available ings. from the immediate region. We therefore Elunoidesforficutus. Swallow-tailed Kite. Rare over measured precipitation totals and daily forested and cleared areas, always in small numbers maximum and minimum temperatures at (a maximum of three birds at once), occurring irregu- Merenberg during most of the study. Rain- larly throughout the year. Buteo mugnirostris. Roadside .* Several pairs fall data for 1976 and temperature data for at forest edge and in cleared areas. Once observed un- the first seven months of that year are pre- successfully attempting to take a small from sented in Table 1. a mixed flock in the forest. In addition to our observations at Finca Miloago chimuchima. Yellow-headed . Common in pastures and at forest edge. Eats large Merenberg, we made brief visits to several numbers of Hercules (Dynustes sp.) in late other upper Magdalena Valley sites. These May and early June. Although primarily a tropical zone include: (1) Puraci: National Park, 2,700- bird, this caracara has taken wide advantage of the new 3,300 m, both slopes Central Cordillera; (2) habitat created by deforestation in the subtropics. Mil- ler (1963) also recorded it at San Antonio in the West- montane forest above San Jo& de , ern Andes, and there is even a specimen from the Bo- 1,900-2,600 m, E slope Central Cordillera; goti plateau, in the temperate zone (Borrero 1952). and (3) Cueva de 10s Guhcharos National F&o .spuruerius. American Kestrel.* Several pairs Park, 1,900-2,000 m, W slope Eastern Cor- resident around cleared areas. Eats many (es- dillera. These visits helped to clarify the pecially orthopterans), also frogs. Copulation observed 28 April; adult (of another pair) seen feeding a fledged distributions of certain species. Figure 1 juvenile 21 May. shows the location of Merenberg, these sub- Ortulis guttutu. Speckled Chachalaca. Common at sidiary sites and Chapmans’ (1917) closest forest edge. Usually seen in pairs, and apparently com- localities. pletely arboreal. Aggressive with Chumuepetes gou- dotii. Ate fruits of Prunus sp. and Cecropiu sp. Loud, STUDY METHODS harsh, repeated calls given regularly in early morning and in evening. This form, columbiunu, approaches its Observations were carried out at Finca Merenherg he- upper elevational limit at Merenherg; it may deserve tween 20 December 1975 and 1 August 1976, except full species status (Meyer de Schauensee 1964). for a lo-day period in mid-June when we visited other Chumuepetes goudotii. Sickle-winged Guan.* Com- sites in the region; RSR was present only during parts mon in forest and at edge. Regularly eats Cecropia sp. of May and June. At Merenherg we cut a l-km trail fruit at dusk at forest edge near the farm dwellings, loop through the forest and designated for study a 1.25- here rather tame and quite habituated to the move- km strip at the forest edge. The forest trail and edge ments of people. Also eats fruits of Ficus spp., Mows strip were both systematically walked at least twice sp., and Turpiniu puniculutu. Did not vary seasonally per week. On encountering a bird we recorded the in numbers. Wing-whirring display was heard quite following data: time of day, its identity, activity, ver- regularly at dusk and before dawn. Although both tical position, and the number and identification of oth- Todd and Carriker (1922) and Miller (1963) suggested er birds with which it was associated (if any). When- that this guan occurs mostly on or near the ground, we, ever the bird was feeding, we attempted to identify the like Johnson and Hilty (1976), found it to be primarily BIRDS OF FINCA MERENBERG 381

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FIGURE 1. Map showing the location of Merenberg, the principal study site, as well as several subsidiary sites and Chapmans’ (1917) nearest localities. 382 ROBERT S. RIDGELY AND STEVEN J. C. GAULIN

arboreal. We saw these guans on the ground near a Pionus tumultuosus. White-capped Parrot.* We pre- small waterfall on several occasions (perhaps drink- fer to follow ONeill’ and Parker (1977) in considering ing?), and on 27 April an adult and two immatures were the northern Andes form, previously given species flushed from the ground near forest edge. rank as P. seniloides, as conspecific with the southern Aramides cujanea. Gray-necked Wood-Rail. Rare on Andes form, nominate P. tumultuosus. An apparent forest floor and at edge, seen only twice, and heard altitudinal migrant from higher elevations; observed only infrequently. Possibly only a wanderer from lower only between 15 July and 1 August (when we departed elevations, whe;e it is numerous, for example, in the the study site), but then common and conspicuous in urmer Magdalena Vallev (Miller 1947). Merenberg an- groups of 10-20. Eats fruits of Turpiniu paniculatu and p&s to ge the highest ‘ location on record foi ’ &e an unidentified member of the Clusiaceae. The two species (Ripley 1977). Merenberg Pionus parrots were never observed to Vunellus chi1ensi.s. Southern Lapwing. At least four flock together. Primarily a parrot of temperate zone for- pairs were resident around wet areas in pastures. A ests, and notably nomadic; Carriker collected four nest with two eggs wars found on 3 January; this failed, higher in Moscopin. but another nest produced two young in February. Amazonu mercenuria. Scaly-naped Amazon.* Com- Tringa solitaria. Solitary Sandpiper. Rare transient. mon in forest; like other parrots, most often observed One &as seen 20 March in a wet part of a pasture. in flight. Seen in pairs or in groups of up to 4-8 birds. Columba fasciutu. Band-tailed Pigeon.* Uncommon Frequently seen flying high overhead in the early in forest canopy and at edge, also perching regularly morning (06:30-08:00), readily identifiable by its large in tall isolated trees. Usually seen in groups of 5-10. size and distinctive shallow wing-beats (like other Its abundance varied greatly, with peak numbers in Amazona), even when color could not be ascertained. early March when Bruneliu comocludifoliu was in When flying in a flock, members of a pair remain close fruit. Also ate fruits of Morus sp. and Ficus cundini- together, one bird remaining behind and slightly to marcensis. This pigeon is probably an altitudinal mi- one side of the other. Ate fruits of Ficus spp., and once grant from higher elevations, but local residents claim seen to eat flesh of baseball-sized fruits of an uniden- that it nests at Merenberg. tified vine. No evident seasonal variation in numbers. Geotrygon montuna. Ruddy Quail-Dove. Very rare Apparently the montane replacement of A. furinosu, on forest floor; possibly overlooked. A single bird was the Mealy Amazon, with similar vocalizations. observed 29 July. Although primarily a bird of tropical Piayu cuyunu. Squirrel Cuckoo.* Rare in forest and zone , this quail-dove has been recorded to 1,900 at edge. Generally more numerous at lower elevations, m in (Meyer de Schauensee and Phelps although Carriker collected three specimens even 1978) and “occasionally” even in the temperate zone higher in Moscop&n. in Colombia (Meyer de Schauensee 1964). Merenberg Crotophugu uni. Smooth-billed Ani. At least one is unquestionably a high elevation for it. group (six individuals) occurred along the edge of a Pyrrhura melanuru. Maroon-tailed Parakeet.* Com- pasture. Merenbergs’ elevation is fairly high for this mon in forest and at edge, frequently perching in iso- species; as Chapman (1917) and others have pointed lated trees. Usually noted in flocks of 8-12 birds, never out, its numbers in the subtropical and lower temper- associating with other parrots. Feeds on fruits of Mi- ate zones increased greatly subsequent to widespread conia theuezuns and other Melastomataceae. Easily the deforestation. Copulation observed 11 June. most numerous parrot at Merenberg. Also common and Ciccaba albitarsus. Rufous-banded Owl. Rare in for- conspicuous at other subtropical sites above the upper est. One was observed for over 2 h on 28 February; Magdalena Valley (e.g., above Isnos and at Cueva de never disturbed, it left its day roost, about 4 m up in 10s G&charos National Park). a Dryopteris tree fern, at 18:lO. Probably overlooked; Hapulopsittucu amazoninn. Rusty-faced Parrot. Very unidentified owl calls were heard occasionally. rare; one group of five seen on 14 June. In flight ap- Streptoprocne zonaris. White-collared . Un- pears obviously smaller than Pionus sp., with shallow- common, in flight over forests and pastures, usually in er wing-beats and a proportionately longer tail. The dispersed flocks. only call heard was an undistinctive, repeated “chek- Phuethornis syrmutophorus. Tawny-bellied . chek, chek-chek .” Not previously recorded from Very rare in forest understory; single birds were seen the upper Magdalena Valley, but G. Tudor (pers. on two occasions, 18 March and 12 May. Mere&erg is comm.) and others observed a probable pair in tem- probably near the species ’ upper altitudinal limit; it perate forest on the E slope of Purac& National Park in was considerably commoner at Cueva de 10s GuCchar- April 1973. The nominate race of this species is known OS National Park. only northward from the Bogota region in the Eastern Colibri delphinue. Brown . Very rare at for- Andes; H. u. fuertisi has not been recorded since the est edge; a single bird netted 15 May. Perhaps the original series was taken in 1911 in highest elevation on record for this widespread but in the northern Central Andes. over 300 km N of Mer- generally rare . enberg (Chapman 1917, Forshaw 1973). Racial alloca- Adelomyiu melunogenys. Speckled Hummingbird.* tion of Merenberg birds is impossible without speci- Abundant in forest understory and undergrowth, to a mens, but the birds seen did show considerable red on lesser extent also at edge. the face, indicating that they were not H. a. fuertisi, Anthocephulu ,fZoriceps. Blossomcrown. Very rare in which has a greenish-yellow face. forest undergrowth; a male seen 19-20 April feeding Pionus chalcopterus. Bronze-winged Parrot. Rare in at a terrestrial orchid (Elleunthus sp.). This observation forest and at forest edge; usually seen only in flight, marks a considerable southward range extension for infrequently perched, and never observed feeding. this little-known hummingbird, in the Central Andes Noted only in May, June, and early July, usually alone previously reported only from northern Tolima De- or in pairs. Calls resemble those of P. menstruws, the partment (Meyer de Schauensee 1966). Carriker, in the well-known Blue-headed Parrot of the lowlands. Prob- Santa Marta Mountains, also found it feeding in forest ably an altitudinal migrant from lower in the subtrop- undergrowth (Todd and Carriker 1922). ical zone: Chapmans’ (1917) collectors obtained a good Urosticte benjamini. Whitetip. Very rare in forest number from Various sites around the head of the Mag- undergrowth; a single immature male of this scarce and dalena Vallev. and K. von Sneidern collected no less little-known hummingbird was netted 15 May. The than 18 spe&ens at La Candela (1,600-1,700 m) in specimen was donated to the Louisiana State Univer- August-September 1942. sity Museum of Zoology (LSUMZ 82900). BIRDS OF FINCA MERENBERG 383

Heliodoxu leadheateri. Violet-fronted Brilliant. gly or in pairs, almost always independent of mixed Common in forest understory, undergrowth, and at flocks. Eats fruits of Ocoteu calophylla and Ficus spp., edge. Although numerous here, Merenberg apparently especially F. cundinimurcensis. Evidently more nu- lies very near the species ’ upper elevational limit; Car- merous at slightly higher elevations in Moscop&n, for riker failed to obtain any, and the species is generally Carriker obtained a series of 10 (and only two P. au- more numerous lower in the subtropical zone (S. Hilty, riceps). pers. comm.). Pharomachrus uuriceps. Golden-headed Quetzal.* Coeligenu coeligenu. Bronzy Inca.* Abundant in for- Rare in middle and upper levels of forest. Behaves est undergrowth and at edge, to a lesser extent also much like P. untisianus, eating the same fruits, but gardens. Nest with two eggs found 19 January inside never seen to feed simultaneously with that species at forest; constructed of mosses, it was 0.7 m above the Merenherg. ground in a crotch of a 1.2-m sapling. The nest failed, Trogon personutus. Masked Trogon. Common in the eggs probably being taken by a squirrel, Sciurus understory and middle levels of forest. Seen singly or granatensis. in pairs, regularly accompanying mixed flocks. Feeds torquatu. Collared Inca.* Abundant in on fruits of vario& Clusiaceae and Piperaceae, and on forest undergrowth, middle levels, at edge, and to a insects, all snatched in flight. Its call is a steady “kwa, lesser extent also gardens. The two incas, so strikingly kwa, kwa, kwa ,” repeated four to eight times, the different in appearance, have quite similar habits; they pitch somewhat higher than the equivalent hut shorter are largely allopatric altitudinally. Chapman (1917) did song of T. collaris, the Collared Trogon. Despite care- note that the two were sympatric at a few Andean lo- ful checking, no T. collaris were noted at Merenberg; calities, but whereas C. coeligena is essentially a snb- that species generally occurs lower in the subtropical tropical species, with Merenberg approaching its up- zone than T. personatus, hut the two are sympatric at per elevational limit, C. torquuta ranges most some localities (e.g., Cneva de 10s Guacharos National commonly in the temperate zone. Higher in Moscopin, Park). Strangely, Carriker obtained no T. per.yonatus Carriker collected 10 C. torquata but only one C. coe- higher in Moscopin; elsewhere it is common well up ligena, whereas, at Cueva de 10s Guicharos National into the temperate zone. Park we found C. coeligenu still to be common and C. Aulucorhynchus prasinus. Emerald Toucanet.* torquata apparently absent. Merenberg seems to lie Common in forest canopy and at edge. Usually seen within the narrow band where both are numerous. singly or in pairs, occasionally in small groups. Fed on Ensiferu ensiferu. Sword-billed Hummingbird.* fruits of Ficus spp., Cecropiu teleulhu, Prunus sp., and Very rare; one netted in a garden 15 May, hut never Sourouhu sp. seen otherwise (although it is a very conspicuous hum- Andigenu nigrirostris. Black-billed Mountain-Tou- mingbird). This individual was presnmably a wanderer can.* Common (though somewhat less numerous than from the temperate zone, where it is not uncommon. Aulacorhynchus) in forest canopy and at edge. Usually We found it on several occasions above Isnos and at observed in groups of two to five birds. Fed on fruits Puraci: National Park, and Carriker collected no less of Ficus spp., Prunus sp., Billia collumbiunu, Cecro- than 10 higher in Moscop6n. pia sp., and Turpinia pan&data. Occasionally ate Boissonneaua jlavescens. Buff-tailed Coronet.* Hercules beetles (Dynastes sp.). Often fed simulta- Common in middle and upper levels of forest and at neously with Chumaepetes and Alouuttu seniculus, edge. Fed at of trees (e.g., Guureu sp.) and at the Red Howler Monkey. A hill rattle was heard more arboreal hromeliads (Guzmania sp.). Holds its wings frequently than the yelping “kree-Cnh, kree-&nh .” up for a second or two after alighting, exposing the call. As Haffer (1974) suspected, this does not prominent rufous under wing coverts; both other cor- seem to occur together with its congener, A. hypog- onets (B. jardini and B. mutthewsi) share this char- laucu, the Gray-breasted Mountain-Toucan, at least in acteristic habit. the Central Andes. Andigena nigrirostris occurs in Heliangelus exortis. Tourmaline Sunangel.* Rare in subtropical and lower temperate forests on both slopes forest understory and at edge. Primarily a temperate (we also found it above Isnos at 2,650 m, and Carriker zone species, mainly above 2,500 m (S. Hilty, pers. collected no less than 12 higher in Moscopin). Andi- comm.). Courtship display observed 7 July: a single genu hypogluucu replaces it in higher temperate for- perched male spread its tail, lowered its head, and ests (3,000-3,300 m). apparently only on the W slope. emitted a series of high rapid chips. Picumnus olivaceus. Olivaceous Picnlet. Very rare Huplophaediu uureliue. Greenish Puffleg.* Uncom- in forest understory and at edge. Found much more mon in forest undergrowth. Noted feeding at flowers commonly in the tropical zone of the upper Magdalena of Pulicourea cuutrecasasii, an understory shrub. At Valley (Miller 1947), and taken by Chapmans’ (1917) San Antonio in the Western Andes, Miller (1963) fonnd collectors as close as La Candela (2,000 m), but rarely it feeding at Palicourea lehmanni. recorded as high as Merenherg. Ocreatus underwoodii. Booted Racket-tail. Very Piculus rivolii. Crimson-mantled Woodpecker.* rare; single males seen on two occasions at forest edge. Common in forest and at edge. Often forages for insects More numerous at lower elevations in the subtropical in epiphytes (e.g., Guzmania sp.); once observed to zone (S. Hilty, pers. comm.; pers. ohserv.). eat the seasonal seeds of Brunelia comocladifoliu. Aglniocercus kingi. Long-tailed Sylph.* Common Melanerpes formicivorus. Acorn Woodpecker. Com- and widespread, from forest (in both nnderstory and mon high in forest, at edge, and in isolated trees. Never canopy) and edge to gardens. In forest, feeds at flowers seen to eat acorns, although these were readily avail- of hromeliads (Guzmania sp.) and vines (Psummisiu able at some seasons; Miller (1963) recorded finding falcuta), among others. Often sallies for flying insects “acorn stores” at San Antonio in the Western Andes. from an exposed canopy perch, especially from 08:OO Occasionally observed sallying after insects from ex- to lo:oo. posed perches. Schistes geoffroyi. Wedge-hilled Hummingbird. Veniliornis fumigutus. Smoky-brown Woodpecker.* Rare in forest understory and at edge. More numerous Rare in forest and at edge. Regularly with mixed flocks, at lower elevations in the subtropical zone, hut never foraging mostly on larger trunks. seems to be a common hummingbird. Veniliornis dignus. Yellow-vented Woodpecker.* Pharomachrus antisiunus. Crested Quetzal.* Rare Uncommon in forest understory, less often at higher (though somewhat more numerous than the next levels (even to the canopy). Frequently with mixed species) in middle and upper levels of forest. Seen sin- flocks, foraging mostly on smaller branches. 384 ROBERT S. RIDGELY AND STEVEN J. C. GAULIN

Phloeoceastes melanoleucos. Crimson-crested ularly associated with mixed flocks. Premnornis is like Woodpecker.* Rare in forest, foraging mostly on large a small foliage-gleaner or Cranioleuca spinetail in its trunks fairly close to the ground. As Meyer de behavior, foraging actively along branches and among Schauensee (1964) pointed out, this essentially tropi- leaves (often dead). Unlike , Premnornis cal-zone woodpecker does occur “occasionally” as rarely if ever actually creeps, nor did we ever see it high as the lower temperate zone. Carriker obtained use its tail as a support. Miller (1963) and S. Hilty (pers. three specimens even higher in Moscopan, and we comm.) have likewise never observed this. Further- once found it at 2,700 m on the E slope of Purace Na- more, the tail of Premnornis lack the promi- tional Park. nent projecting shafts of Premnoplex; Premnornis is Xiphocolaptes promeropirhynchus. Strong-billed not a “barbtail” at all. As the two are so different in .* Uncommon in forest and at edge, for- behavior, we believe Vauries’ (1971) merging of the aging mostly on larger trunks and branches at all levels. genera Premnornis and Premnoplex into Usually met with singly, and not with mixed flocks. is questionable. Premnoplex does seem close to Mar- Generallv dendrocolantid-like in its foraging stvle. but garornis (which acts much like a woodcreeper), but also seen to splinter dead wood. Its unmistakable ’ call Premnornis certainly does not seem similar. is a whistle-like series of four to five pairs of descend- Pseudocolaptes boissoneautii. Streaked Tufted- ing notes, the first of each pair being the same pitch or cheek.* Uncommon in upper and middle levels of for- slightly lower than the second of the preceding pair, est, occasionally coming lower. Frequently with mixed the whole often initiated with a single higher note. flocks. Forages especially in large bromeliads, some- This call is given almost exclusively at dawn and dusk, times seeming to almost disappear inside them. and is delivered at very long intervals. Three other Syndactyla subalaris. Lineated Foliage-Gleaner. members ofthe (X. albicollis, X.faZcirostris, and Uncommon in forest, foraging at all levels, but least X. major) all give virtually identical calls. often in the canopy. Regularly with mixed flocks. Xiphorhynchus triangularis. Olive-backed Wood- Anabacerthia striaticollis. Montane Foliage-Glean- creeper.* Uncommon at middle levels of forest, less er. Common at all levels of forest and at edge. Usually often at edge. Only an occasional member of mixed seen with mixed flocks. Carrikers’ failure to collect flocks, where its presence was positively correlated either Syndactyla or Anabacerthia higher in Mosco- with that of Syndactyla subalaris. pin would seem to indicate that Merenberg must be Lepidocolaptes affinis. Spot-crowned Woodcreeper.* near the upper altitudinal limit for both species. Common at all levels of forest and at edge. A frequent rutilans. Streaked Xenops.* Rare at middle member of mixed species flocks, where its presence and upper levels of forest and at edge. Usually seen was positively correlated with that of Anabacerthia with mixed flocks. Forages mostly on smaller branches striaticollis. Aggressive with Xiphorhynchus triangu- and twigs, gleaning actively like a chickadee (Parus Zaris. sp.). More numerous lower in the subtropical zone, Synallaxis azarae. Azaras’ Spinetail.* Abundant in though Carriker did collect three even higher in Mos- dense vine tangles and in forest undergrowth, espe- cop&n. cially at openings and edge. Usually in pairs, and not unicolor. Uniform Antshrike.* Very moving with mixed flocks. Not easy to see, but its rare; a male mist-netted 14 May. Though termed in the abundance is revealed by its constantly reiterated “ka- literature as an “upper tropical and lower subtropical” keek, ka-keek ..” species (Meyer de Schauensee 1950, 1964, 1970), Car- Cranioleuca curtata. Ash-browed Spinetail. Very riker obtained four even higher in Moscopin (2,500 m). rare in forest understory and at middle levels; a single This is virtually in the lower temperate zone and prob- individual 12 July. Somewhat more numerous at Cueva ably the highest the species has ever been found. This de 10s Gu6charos National Park; Merenberg is proba- antshrike seems everywhere to be a scarce and incon- bly near the upper elevational limit for the species. A spicuous species (S. Hilty, pers. comm.; pers. observ.). rare bird in Colombia. Cranioleuca curtata behaves Terenura caZZinota. Rufous-rumped Antwren. Un- much like the better-known C. erythrops, the Red- common in middle and upper levels of forest. Forages faced Spinetail; it is arboreal, gleaning actively and actively with mixed flocks, often exposing its rufous even acrobatically in foliage and among epiphytes, es- rump as it peers under branches or briefly hangs up- pecially on larger branches. It is frequently found with side-down. Merenberg appears to be the highest lo- mixed flocks. cality on record for the species (Meyer de Schauensee Siptornis striaticollis. Spectacled Prickletail. Very 1950, Meyer de Schauensee and Phelps 1978). Its rare in forest understory, on lower branches of larger small size and habit of foraging well above the ground forest trees, and at edge; seen 27 February and 28 June. probably account for its relative rarity in collections; Somewhat more numerous at Cueva de 10s Guicharos locally it can actually be fairly common. National Park. Rare in collections, previously recorded Grullaria ruficapilla. Chestnut-crowned Antpitta. in Colombia only from 1,650 to 2,000 m (Meyer de Common on or near the ground in forest and at edge, Schauensee 1950), and so far only from the slopes especially in the dense growth which succeeds tree- above the Magdalena Vallev. Merenberg is the highest falls. Very difficult to observe, although its nasal whis- locality on record for the species. Siptornis was-seen tled call is often heard; the bird will decoy to a good singly, invariably with mixed flocks; it foraged much whistled imitation of it. The call consists of three notes, like a Xenops, working along outer branches and on the first sometimes inaudible or omitted, the second twigs, occasionally even hanging upside-down like a lower, the third rising; it is given four to seven times chickadee (Parus sp.). We never heard any vocaliza- a minute. We found another antpitta, the Bay-backed tion. Antpitta (G. hypoleuca), to be common in dense forest Premnoplex brunnescens. Spotted Barbtail. Uncom- edge and in overgrown clearings at Cueva de 10s GuC- mon in forest undergrowth and understory. As Miller charos National Park (1,900-2,000 m). This species has (1963) and others have noted, this species forages by been recorded as close to Merenberg as La Candela, creeping over branches and trunks, sometimes using at 1,950 m (Meyer de Schauensee 1950), but we never its tail for support as do the . Sometimes heard its distinctive three-noted call (a first long one associated with mixed flocks of undergrowth birds, but followed by two unslurred shorter and higher ones) more often occurred alone. here. Merenberg probably is a little too high for this Premnornis guttuligera. Rusty-winged Barbtail.* upper tropical and subtropical antpitta. Common in forest undergrowth and understory. Reg- sp. .* Although we never saw a BIRDS OF FINCA MERENBERG 385 tapaculo, we occasionally heard the characteristic vo- Leptopogon); rather active, and, unlike the preceding calization of some member of this genus coming from small flycatcher, not at all hard to see. dense forest undergrowth. Carriker collected three Mecocerculus poecilocercus. White-tailed Tyrannu- specimens of S. unicolor, the Unicolored Tapaculo, let. Uncommon in upper and middle levels of forest. higher in Moscop&n, but two other species (S. femor- Seen singly or in pairs, frequently with mixed flocks. alis and S. latebricola) are also possible here. Often exposes white in tail. Regularly gives a distinc- Pipreola riefferii. Green-and-Black Fruiteater.* tive high sibilant “tee-tee-tee-tee.” Its horizontal war- Common in understory and middle levels of forest. hler-like posture is strikingly unlike the upright pos- Seen singly or in pairs, sometimes with mixed flocks, ture of M. Zeucophrys, the White-throated Tyrannulet, invariably approachable and lethargic. Eats fruits of found at higher elevations. understory trees and vines, notably Hedyosum sp. Eluenia sp. Elaenia.* Rare at forest edge. Carriker Pachyramphus uersicolor. Barred Becard.* Common collected four specimens of E. pallntungue, the Sierran in upper and middle levels of forest and at edge. A Elaenia, higher in MoscopCn, and as he obtained no frequent member of mixed flocks; the males’ pretty other Elaeniu, Merenberg birds were probably this whistled “treedididee?” was often heard. Although species. numerous and conspicuous here, Carriker obtained Tyrnnniscus uiridijuuus. Golden-faced Tyrannulet.* only one at higher elevations in Moscopin, indicating Uncommon at forest edge and in middle and upper that this becard is approaching its upper elevational levels of forest. Most often with mixed flocks. Perches limit at Merenberg. horizontally, often with its tail “half-cocked.” Ochthoeca cinnamomeiuentris. Slaty-backed Chat- Leptopogon rufipectus. Rufous-breasted Flycatch- Tyrant.* One pair observed throughout the study in an er.* Common in understory and middle levels of forest. area of dense forest undergrowth near a small waterfall. Usually seen singly; sometimes with flocks. Superfi- Both members of the pair were observed building a cially resembles Myiophobus pulcher, and best distin- nest of mosses on a rock ledge behind the waterfall on guished by its larger size, grayish crown, and rufous 15 February. facial area, also by its frequent habit of lifting its wings, Tyrannus melancholicus. Tropical .* Un- one at a time, up over its back, as do all members of common around isolated trees and at edge of pastures. the genus (Ridgely 1976, pers. ohserv. for L. tucza- Much more numerous at lower elevations (e.g., in the nowskii). Probably here approaching its upper eleva- upper Magdalena Valley; Miller 1947), but also regu- tional limit; Carriker obtained only one specimen larly recorded up into the temperate zone in cleared higher in Moscopin. areas (Meyer de Schauensee 1950, pers. observ.). Car- Mionectes striaticollis. Streak-necked Flycatcher.* riker obtained one even higher in Moscop&n. Rare in forest understory and at edge. Usually seen Myiodynustes chrysocephalus. Golden-crowned singly, often with mixed flocks. Flycatcher. Very rare; one observed at forest edge on Notiochelidon cyanoleuca. Blue-and white Swal- 21 April. Probably a wanderer from lower in the suh- low.* Abundant over cleared areas, especially around tropical zone; Merenbergs’ elevation is relatively high buildings, which provided nest sites. for the species (Chapman 1917, Meyer de Schauensee Cyunolyca viridicyunu. Collared Jay. Very rare; a 1950). group of three or four birds seen only l-2 June. The Contopus fumigutus. Greater Pewee.* Very rare at group foraged independently of other birds in middle forest edge; one seen on 7 February. A widespread and and upper levels of forest. Collared Jays were also seen usually numerous bird of the subtropical and temper- in forest above Isnos (2,500 m), but judging from the ate zones (Meyer de Schauensee 1950, pers. observ.). few records mentioned by Meyer de Schauensee We found it common at elevations both lower (Cueva (1951), the species is not numerous around the head of de 10s GuLcharos National Park) and the same (ridge the Magdalena Valley. above Isnos) as Merenberg, and Carriker collected 10 Cyanocorur ynccls. Green Jay. Common and con- higher in MoscopLn. Its rarity at Merenberg is inexpli- spicuous in all levels of forest, at edge, and in isolated cable. trees. Forages actively and noisily in groups of two to Pyrrhomias cinnamomea. Cinnamon Flycatcher.* six, usually not associating with other birds. Unlike Abundant at edge, much less numerous at middle what Alvarez (1975) found in a more disturbed area in levels in forest, then usually at openings. Most often northern Colombia, Green Jays at Merenberg were not seen in pairs, usually not with flocks. Perches very up- especially shy, and indeed often seemed inquisitive, right, sallying out to capture insects from the air and sometimes closely approaching the observer, especial- from foliage. ly in response to squeaking. Their loud discordant call Myiophobus fZauicuns. .* notes are characteristically tripled (e.g., “renh-renh- Common at edge and in middle and understory levels renh”) in quality and rhythm remarkably reminiscent of forest. Seen in twos and threes, rarely with flocks. of a North American katydid (Pterophyllu sp.)! Carri- Typically forages by sallying out from a low perch, kers’ not obtaining any higher in MoscopCn would in- picking off insects from leaves, branches, or the dicate that Merenberg is near the species ’ upper ele- ground. Its posture is upright; the bird recalls an Em- vational limit. pidonax. Odontorchilus branickii. Gray-mantled . Rare Myiophohus p&her. Handsome Flycatcher.* Com- in upper and middle levels of forest. Forages among mon at middle and upper levels of forest, less often at mosses and other epiphytes on larger branches, gen- edge. Seen in small groups of two to four birds accom- erally avoiding the leafy tips. Seen with mixed flocks, panying large mixed flocks. In appearance strikingly usually large. A very distinctive wren, quite gnatcatch- like the larger Leptopogon ru&pectus (see below). er-like in its plumage, foraging style, and attitudes Poecilotriccus rujiceps. Rufous-crowned Tody-Ty- (even to the often-raised tail). Strikingly quiet for a rant.* Uncommon in dense undergrowth at edge of for- wren; we have not distinguished its calls anywhere in est and in openings. Seen singly, not with flocks. its range. This race (nominate 0. h. hrunickii) has pre- Perches in low shrubbery, making short abrupt sallies to glean insects off foliage. viously been recorded in Colombia from only one Pogonotriccus ophthalmicus. Marble-faced Bristle- specimen, that taken at La Palma, 1,650 m (Chapman Tyrant. Rare in middle and understory levels of forest. 1917, Meyer de Schauensee 1951). Hilty (1977) re- Regularly joins mixed flocks. Perches upright, occa- cently found the western race (minor) in the upper sionally flicking a wing up over its back (much like tropical zone on the W slope of the Western Andes in 386 ROBERT S. RIDGELY AND STEVEN J. C. GAULIN

Valle Department. Mere&erg is the highest locality at edge. Usually foraged with mixed flocks of resident on record for the species. insectivores. Last seen 10 May. Thryothorus genibarbis. Moustached Wren. Com- Parulu pitiayumi. Tropical Parula. Uncommon in mon in dense undergrowth in forest and at small open- middle and upper levels of forest and at edge. Fre- ings. Very difficult to observe, hut its rich musical quently forages with mixed flocks, usually in pairs. phrased song was often heard emanating from impen- Dendroica fuscu. Blackburnian Warbler. Abundant etrable tangles. Merenbergs’ elevation is probably near winter resident in middle and upper levels of forest the upper limit for the species; Carriker did not obtain and at edge. The most commonly observed bird during any. the northern winter, as it was near Popay6n, Cauca Troglodytes aedon. House Wren. Uncommon in Department, in disturbed oak woodland at 1,650 m brush along pasture edges. (Chipley 1976). It was also the most numerous North Troglodytes solstitialis. Mountain Wren.* Common American migrant at San Antonio in the Western Andes in vine tangles and in undergrowth of forest. Some- (Miller 1963). A very frequent member of mixed times forages with mixed flocks, hut also often alone. species flocks. Last seen 24 April; Miller (1963) last On 22 March observed bringing nesting material, fine recorded it on 20 April, but we also saw a female along epiphyte rootlets, to a cluster of dead Cecropiu leaves the old Buenaventura road (600 m on W slope of West- about 7 m above the ground at forest edge. ern Andes in Valle Department) on 5 May. Henicorhina leucophrys. Gray-breasted Wood-Wren.* Wilsoniu canadensis. Canada Warbler.* Common Uncommon in dense forest undergrowth, principally winter resident, foraging in vine tangles and forest un- around overgrown treefalls. More numerous lower in dergrowth. Chipley (1976) and Miller (1963) also found the subtropical zone, e.g., at Cueva de 10s Guhcharos this species the second most numerous migrant. Often National Park (where it was abundant). forages with mixed species flocks, especially with Bus- Mimus giluus. Tropical Mockingbird. Rare around ileuterus spp. Last observed 8 April; Miller (1963) last isolated trees and along fence rows in pastures. More recorded it on 13 April, hut we also saw a female along numerous at lower elevations in the upper Magdalena the old Buenaventura road (500 m on W slope of West- Valley (Miller 1947, pers. observ.). ern Andes in Valle Department) on 5 May. Myadestes ralloides. Andean Solitaire.* Uncommon Myioborus miniatus. Slate-throated Redstart.* at forest edge, and in forest understory. Eats fruits of Abundant, foraging very actively in more diverse mi- Hedyosmum sp. crohabitats of forest than any other Merenberg bird: in Turdus j&cater. Great .* Rare at forest edge understory, middle and upper levels, vine tangles, and in pastures. Eats fruits of Cecropia sp. Much more edge, even on trunks and large horizontal branches. numerous at higher elevations, but occurs widely in Frequently makes short sallies, catching insects in the cleared parts of the subtropical zone (Meyer de air, apparently having first flushed them. Often forages Schauensee 1951, pers. observ.). with mixed flocks, but is also regularly seen alone or Turdus serranus. Glossy-black Thrush.* Rare at for- in pairs. est edge and in forest. Eats fruit of Cecropia sp. Much Basileuterus Zuteooiridis. Citrine Warbler. Very rare less conspicuous than the preceding species. More nu- at forest edge; one seen with a mixed flock 14 Febru- merous in forest above Isnos, at slightly higher (2,400 ary. More common above Isnos (2,400-2,500 m), where m) elevations than Merenberg; Carriker collected four it also fed at edge. higher in MoscopCn. Busileuterus tristriutus. Three-striped Warbler. Turdus ignobilis. Black-hilled Thrush. Rare at forest Common in forest undergrowth and around overgrown edge. Much more numerous at-lower elevation in semi- treefalls. Usually in small groups, less often in pairs; open areas in the Magdalena Valley (Miller 1947, pers. frequently forages with mixed flocks. Carriker failed to ohserv.). obtain any higher in MoscopLn, indicating that despite Cyclarhis nigrirostris. Black-hilled Peppershrike.* its abundance here, Merenherg must be near its upper Uncommon in middle and upper levels of forest and elevational limit. at edge. Cyclarhis gujanensis, the Rufous-browed Busileuterus coronutus. Russet-crowned Warbler.* Peppershrike, replaces nigrirostris at lower elevations Abundant in forest undergrowth and around over- in the upper Magdalena valley; we found it as high as grown treefalls, sometimes feeding on the ground in near La Plata (1,300 m) and Palestina (1,450 m). Cy- leaf litter and at edge. Often with mixed flocks, fre- cZarhis nigrirostris behaves and sings very much like quently associating with B. tristriatus. Seen feeding the better-known C. gujunensis, but seems to require a fledged juvenile 6 June. more extensive forest, and does not tolerate mostly Conirostrum albifrons. Capped Conehill.* Common cleared areas (in which C. gujunensis often occurs). at upper levels of forest and at edge, where it some- Vireo gilwus. Warbling Vireo.* Abundant in middle times comes lower. Most often forages by gleaning in- and upper levels of forest and at edge. A frequent sects from terminal leaves and branches, but also takes member of mixed flocks. some Cecropia sp. fruit. Often wags tail. Usually seen Vireo olivuceus. Red-eyed Vireo. Rare winter resi- with mixed flocks. dent, noted most often in middle levels of forest. Did Diglossu ulbilatera. White-sided -piercer.* not sing. Last seen 26 March. Uncommon in forest understory and at middle levels, lcterus chrysuter. Yellow-backed Oriole.* Uncom- also at edge and in gardens. Seldom with mixed flocks. mon at forest edge and in isolated trees in pastures. Regularly observed piercing flowers, especially those More numerous at lower elevations in the upper Mag- of Palicourea cuutrecasusii, but also gleans insects dalena Valley (Miller 1947, pers. observ.), but Carriker from foliage. did obtain two specimens even higher in Moscopin Diglossa cyuneu. Masked Flower-piercer.* Rare in (which must he nearly the highest elevation ever re- middle levels of forest, in understory, and at edge. Much more numerous in temperate zone forests, and corded for this species). common in the Puraci: National Park area; Carriker col- SturneZZa magna. Eastern Meadowlark.* Rare in pas- lected eight higher in Moscopin. tures. More numerous at lower elevations in the upper Euphonia xunthoguster. Orange-bellied Euphonia.* Magdalena Valley (pers. ohserv.); Carriker, however, Rare in middle and upper levels of forest and at edge. obtained one specimen even higher in Moscopin. More numerous lower in the subtropical zone (e.g., at Vermioora peregrinu. Tennessee Warbler. Rare win- Cueva de 10s Guicharos National Park), hut Carriker ter resident in middle and upper levels of forest and did obtain one even higher in Moscopin. BIRDS OF FINCA MERENBERG 387

Pipraeidea melanonota. Fawn-breasted . pecially Cecropiu spp. Occasionally gleans insects Uncommon at forest edge and in isolated trees in pas- from leaves. tures. Thraupis episcopus. Blue-gray Tanager. Rare in iso- Tungaru urthus. Golden Tanager.* Rare in forest and lated trees in pastures and at forest edge. Much more at edge. Seems notably less numerous in the upper numerous at lower elevations in the upper Magdalena Magdalena Valley than in the Western Andes (pers. Valley (Miller 1947, pers. ohserv.). ohserv.). Merenhergs’ elevation is rather high for this Pirungu ruhru. Summer Tanager.* Rare winter resi- essentially subtropical species; Carrikers’ single spec- dent; last observed 7 March. Forages at edge and in imen from even higher in Moscopin is among the high- middle and upper levels of forest; noted singly, usually est ever recorded for the species. Frequently forages with mixed flocks. on bare or mossy branches (even trunks) in a manner Tachyphonus rufus. White-lined Tanager. Very rare much like that of the Black-and-white Warbler (Mni- at forest edge; one seen 22 May. Considerably more otilta uariu). Willis (1966) and Hilty and Simon (1977), numerous lower in the Magdalena Valley. in the Western Andes, noted that several Tangaras em- Creurgops uerticulis. Rufous-crested Tanager.* ployed this method of searching for insects; we found Common in upper levels of forest and at edge. Never that among Merenherg members of the genus, T. ar- seen to eat fruit; gleans insects, almost always in leafy thus employed this method most frequently. terminal areas. Also flycatches opportunistically, and Tangara xanthocephalu. Saffron-crowned Tanager.* much more frequently than any other Merenherg tan- Abundant in middle and upper levels of forest and at ager. Usually accompanies mixed flocks. edge. More frugivorous than any other resident Tan- Chlorospingus ophthalmicus. Common Bush-Tana- guru, eating especially Cecropia “pp., Morus sp., Mi- ger.* Abundant in-upper and middle levels of forest, conzu spp., and other members of the Melastomata- in understory, and at edge. Usually in groups, most ceae. Less often gleaned for insects in leafy terminal often with mixed flocks. areas. Cnemoscopus rubrirostris. Gray-hooded Bush-Tan- Tungaru chrysotis. Golden-eared Tanager. Uncom- ager.* Very rare at middle levels of forest and at edge; mon in middle and upper levels of forest and at edge. seen 21 January and 5-9 June. Seen singly or in pairs, As only one specimen of T. chrysotis has been record- usually with mixed flocks. Often wags tail. More nu- ed from Colombia, that taken at San Agustin (Meyer merous at higher elevations (e.g., ahove Isnos and in de Schauensee 1951), we were surprised to find this Puracir National Park); Carriker took four higher in species regularly, in small numbers, at Merenherg. We Moscopin. also saw a few individuals along the trail from Palestina Hemispingus frontalis. Oleaginous Hemispingus. to Cueva de 10s Guicharos National Park (1,850 m). Common in dense forest undergrowth and in vine tan- Usually observed eating melastome fruit; also seen eat- gles. Often accompanies mixed flocks of Busileuterus ing fruits of Morus sp. and gleaning insects from spp. and Premnornis. Carriker failed to obtain any leaves. higher in Moscopin, indicating that Merenherg is near Tanguru parzudakii. Flame-faced Tanager.* Com- the species ’ upper elevational limit. mon at edge and in upper levels of forest. Forages for Hemispingus melunotis. Black-eared Hemispingus. insects almost exclusively in lichens (Usnea sp.) epi- Very rare in forest undergrowth; seen 6 and 28 July. phytic on tree branches; only rarely gleans in leafy tips. Seen in pairs or singly, often foraging with mixed Tunguru parzudukii also eats fruit of Cecropia spp., flocks. A subtropical tanager, rather local, generally as- hut not nearly as frequently as T. xunthocephalu. sociating with stands of bamboo Chusqueu sp.; not Courtship feeding observed 21 February. previously reported between the Department of Toli- Tanguru labradorides. Metallic-green Tanager.* ma in the Central Andes and Nariiio Department (Mey- Rare at forest edge. Observed feeding on Cecropiu er de Schauensee 1951, 1964). J. Silliman also took spp. fruit, and also gleaned in leafy terminal branches. several specimens in 1971-1972 on the E slope of Pur- More numerous lower in the subtropical zone, espe- a& National Park; these now reside in the INDER- cially in the Western Andes (Miller 1963, pers. oh- ENA collection in Bogoti (S. Hilty, pers. comm.). serv.). Carriker obtained one specimen in MoscopCn; Atlupetes gutturalis. Yellow-throated Brush-Finch. this is prohahly the highest elevation ever recorded for Rare at edge and around overgrown treefalls in forest. the species (Meyer de Schauensee 1951). Atlupetes fuscooliuuceus. Dusky-headed Brush- Tanguru ruficeruir. Golden-naped Tanager. Com- Finch.* Rare in forest undergrowth, especially around mon in upper and middle levels of forest and at edge. overgrown treefalls and at edge. This upper Magdalena Eats fruits of Cecropiu spp. and Mot-us sp., as well as Valley endemic is more numerous at lower elevations gleans insects from leaves. Merenherg appears to he in the subtropical zone, e.g. at San Agustin and Cueva one of the highest recorded localities for this typically de 10s Guacharos National Park. Merenherg ohserva- subtropical species (Meyer de Schauensee 1951); Car- tions and Carrikers’ pair of specimens considerably riker failed to obtain any higher in Moscopan. extend the upper elevational limit of this species, pre- Tangura nigrouiridis. Beryl-spangled Tanager.* viously recorded only at 1,550-1,900 m (Meyer de Common at middle levels of forest and at edge. Except Schauensee 1951). The seemingly closely related A. for T. xunthocephalu, this species was the most frugi- fluuiceps, the Olive-headed Brush-Finch, was recently vorous Tunguru. Frequently associates with the latter, mist-netted and photographed by J. S. Dunning not far particularly at fruiting melastomes and Mows sp. from Merenherg; the species was previously known When T. nigroviridis forages for insects, it often only from northern , well to the searches hare terminal branches. north of our area (Meyer de Schauensee 1951, 1970). Tanguru uussorii. Blue-and-Black Tanager.* A proh- Three birds were captured on 18-19 December 1967 able altitudinal migrant; observed only from 25 De- near La Plats Vieja, ca. 1,300 m, further down the Rio cember to 2 January, then feeding especially on sea- de la Plats Valley. We examined the photographs and sonal fruits of Miconiu sp. Much more numerous in the found the birds clearly referable to this species. Atlu- temperate zone; Carriker obtained a series of 11 higher petes flaoiceps here appears to he virtually sympatric in Moscopin. with A. fuscooliuuceus, perhaps replacing that species Anisognuthus jhzoinuchu. Blue-winged Mountain- Tanager.* Abundant at edge and in middle and upper lower in the subtropical zone. levels of forest. Highly frugivorous, exploiting Prunus Atlapetes brunneinuchu. Chestnut-capped Brush- sp., Miconiu sp., Morus sp., Souroubeu sp., and es- Finch.* Rare in forest undergrowth, often foraging on 388 ROBERT S. RIDGELY AND STEVEN J. C. GAULIN

the ground. Perhaps somewhat overlooked. Usually in ably disappeared mostly as a result of the extensive pairs, and not associating with mixed flocks. deforestation that has taken place in recent decades, Zonotrichia cupensis. Rufous-collared Sparrow.* perhaps also due to some direct persecution. It has also Common in pastures and at edge. declined elsewhere in its rather limited range, and has Spinus psaltriu. Dark-backed Goldfinch. Very rare; been given “vulnerable” status in the latest Red Data one male mist-netted 14 May. More numerous lower Book (King 1979). Yellow-eared Parrots presumably in the Magdalena Valley. used to occur, at least seasonally, at Merenberg, but it is doubtful that they ever will again. CARRIKERS’ COLLECTIONS FROM Tyto u&u. Barn Owl. Local in clearings and open MOSCOPAN areas virtually throughout Colombia (Meyer de Schauensee 1964), we never found Barn Owls at Mer- M. A. Carriker, Jr., a professional collector enberg, but they could occtlr. then residing in Colombia, made at least Colibri thulussinus. Green Violetear. Although this three major collecting trips to Moscophn af- conspicuous hummingbird appears to be common only ter the Popaybn-Neiva highway was opened a little higher than Merenberg, and also occurs regu- larly at certain localities lower in the subtropical zone to traffic in the late 1940s: February-March (Meyer de Schauensee 1964), we never recorded it at 1952, March 1954, and February 1956. Dur- Merenberg. It could occur sporadically here. ing these periods, Carriker obtained a total Lufresnuyu Zufresnuyi. Mountain Velvetbreast. In of 501 specimens representing 114 species. this part of Colombia Lufresnuya appears to be re- stricted to the temperate zone, where it occurs in Only a very few of these specimens have shrubby areas and at forest edge (S. Hilty, pers. comm.; been recorded in the literature. Judging pers. observ.). It almost certainly never occurs as low from his specimen labels, most of his col- as Merenberg. lecting was carried out near an elevation of Eriocnemis alinue. Emerald-bellied Puffleg. Chiefly 2,485 m, or about 185 m above Merenbergs’ a subtropical zone species inhabiting the interior forest (Meyer de Schauensee 1970, pers. observ.), this puf- average elevation. Some of his specimens, flea, although generally an uncommon hummingbird, of course, doubtless came from somewhat seems likely to occur at Merenberg. Possibly it was above or below this elevation. overlooked. Carrikers’ Moscophn birds help put the Ramphastos umhiguus. Black-mandibled Toucan. Moscopin is the highest locality ever recorded for this Merenberg avifauna into better perspective, toucan (Haffer 1974); Carrikers’ specimen may have especially coming as they do from a slightly been a wanderer from lower in the subtropics. None higher elevation. Many species are of was ever recorded at Merenberg, nor was the species course found in both areas; we have com- (an obvious one) known to local residents. Apparently mented above on a number of these. Below more numerous, though now far from common, lower in the subtropical zone around the head of the Mag- we briefly discuss only the 34 species which dalena Valley; Chapmans’ (1917) collectors obtained Carriker collected but which we did not several from various sites at elevations ranging from find at Merenberg. 1,500 to 2,000 m. Forests at these levels, it should be noted, have now been extensively cleared. Nothocercus julius. Tawny-breasted Tinamou. No tin- Cumpylorhumphus pucherunii. Greater Scythebill. amou was seen or heard at Merenberg. This one, wide- Apart from “BogotL” trade-skins (with no precise data), spread but elusive in temperate zone forests (or per- Carrikers’ specimen appears to be the first record of haps N. bonupartei, the Highland Tinamou, which re- this rare and poorly-known woodcreeper in Colombia places it at subtropical elevations), might formerly have away from the Western Andes (Meyer de Schauensee been found here, but if so it seems to have been locally 1950). extirpated. Murgarornis squumiger. . A Mergunetta armuta. Torrent Duck. Merenberg lacks common bird of temperate zone forests, numerous at the swift rocky streams or rivers that this duck requires. Purac6 National Park, it almost certainly never occurs Buteo plutypterus. Broad-winged Hawk. Probably as low as Merenberg. occurs in small numbers at Merenberg, as a transient Ampelion rubrocristatus. Red-crested . or winter visitant from North America. An unidentified Found chiefly in the temperate zone up to the edge of Buteo sp. (not B. magnirostris) seen 7 March may well pkramo, this cotinga would not be expected to occur as have been this species. low as Merenberg. Herpetotheres cuchinnnns. Laughing Falcon. Pri- Ampelion rufuxillu. Chestnut-crested Cotinga. Un- marily a tropical zone raptor (Brown and Amadon like the previous species, A. rufuxillu is a bird of sub- 1968), Herpetotheres seems to have spread upward tropical forests, where it perches conspicuously in the into the subtropical zone subsequent to local defores- canopy and often sallies after insects. Given Carrikers’ tation. Moscopin may be the highest locality ever re- four specimens from only a little higher in Moscopin, corded for the species. We also saw a pair above Isnos its absence at Merenberg is puzzling, but could be re- (at 2,300 m), but it was never recorded at Merenberg. lated to the small extent and relative isolation of Mer- Ognorhynchus icterotis. Yellow-eared Parrot. Ap- enberg forests, in conjunction with the species ’ relative parently this parrot was quite frequent in the Mosco- rarity throughout its range. p&n region in the 1950s. In addition to the five speci- Lipuugus fuscocinereus. Dusky Piha. Presumably a mens obtained by Carriker, F. C. Lehmann (1957) very vocal bird (as is its close relative of the Amazonian collected a number of others in the same general area. lowlands, L. uociferuns, the Screaming Piha), this However, Lehmann (1957) was already noticing a species could not have been overlooked if it had been steady diminution in numbers, and this decline has present at Merenberg. It seems to occur chiefly in the now continued to the point where this species has temperate zone (Meyer de Schauensee 1950), which largely disappeared from the region. Except for a pair perhaps helps to account for its absence here. Perhaps, in flight on one occasion over the pCramo at Purack also, the remaining patches of forest are too small for National Park, we never recorded it. This parrot prob- it. BIRDS OF FINCA MERENBERG 389

Pyroderus scututus. Red-ruffed Fruitcrow. This numerous in Pura& National Park. As Carriker ob- large cotinga has seemingly been extirpated from the tained six, and as we also observed it as low as 2,400 immediate Merenberg area. m above Isnos, it seems possible that this tanager, like Rzlpicola peruuiana. Andean Cock-of-the-Rock. Like Tanguru vussorii, might occasionally wander as low as the fruitcrow, almost certainly extirpated from Mer- Merenberg. enberg; local residents told us that it used to occur. Buthruupis montana. Hooded Mountain-Tanager. The streamside habitat that the species favors is not Found chiefly in temperate zone forests, although re- found at Merenberg, so, presumably, it never was pres- corded somewhat lower in the Western Andes (Chap- ent in large numbers. man 1917, Meyer de Schauensee 1951), this tanager Ochthoeca fumicolor. Brown-backed Chat-Tyrant. probably never occurs as low as Merenberg. Common in temperate zone clearings up to the edge Dubusia taeniutu. Buff-breasted Mountain-Tanager. of piramo (Meyer de Schauensee 1964, pers. observ.), Occurring mostly in temperate zone clearings and at Moscopin is unusually low for the species (and is near- forest edge (Chapman 1917, pers. observ.), this tanager ly the lowest ever recorded for it). Ochthoeca fumi- could occasionally wander as low as Merenberg, for color would certainly never occur as low as Meren- Meyer de Schauensee (1951) recorded it locally from berg. the subtropical zone as well. Myiarchus cephalotes. Pale-edged Flycatcher. We Chlorornis riefferii. Grass-Green Tanager. Evident- never saw a Myiarchus at Merenberg. Myiarchus ly quite numerous only a little higher than Merenberg cephulotes is recorded widely from the subtropical (Carriker obtained seven in Moscop&n), and seen by zone around the head of the Magdalena Valley, mostly us as low as 2,400 m above Isnos, this tanager may at somewhat lower elevations than Merenberg (Meyer occur at least occasionally at Merenberg itself. Al- de Schauensee 1950). Its absence from Merenberg is though commonest in the temperate zone, Chlorornis puzzling. has also been recorded locally from the subtropics, e.g., ldioptilon granudense. Black-throated Tody-Tyrant. at La Candela (2,000 m) in the upper Magdalena Valley Though fairly conspicuous and, unlike so many small (Meyer de Schauensee 1951). Andean flycatchers, easy to identify, we failed to find Sporophilu nigricollis. Yellow-bellied Seedeater. this tody-tyrant at Merenberg. It might be expected to Although essentially a tropical zone bird, this seedeat- occur, for we did observe it above Isnos (2,400 m) and er has also been recorded locally in the subtropics at Cueva de 10s Gu&charos National Park (1,900 m), (Meyer de Schauensee 1951). Like Spinus psultria, it and also found it quite common in temperate forests in may occur as an occasional wanderer at Merenberg. Purack National Park. Its apparent absence from Mer- Atlupetes schistuceus. Slaty Brush-Finch. Although enberg is difficult to explain. common at elevations almost equal to Merenbergs’ on Pseudotriccus rujiceps. Rufous-headed Pygmy-Ty- the ridge above Isnos (2,400 m up), this conspicuous rant. A shy bird of forest undergrowth, this flycatcher and rather arboreal brush-finch is evidently scarce in could have been overlooked at Merenberg. In our ex- Moscopin, as Carriker collected only one. It apparent- perience, however, it occurs chiefly in the temperate ly does not occur at Merenberg. zone, somewhat lower in the Western Andes (S. Hilty, pers. comm.; pers. observ.). DISCUSSION Pogonotriccus poecilotis. Variegated Bristle-Tyrant. Never certainly identified at Merenberg, this flycatch- During our nine months of study at Finca er is recorded from a number of lower subtropical lo- Merenberg, we recorded 132 bird species. calities in the upper Magdalena Valley (Meyer de As discussed above, Carriker obtained a fur- Schauensee 1950). It probably does occur in small ther 34 species in Moscophn during three numbers at Merenberg. lengthy collecting trips in the 195Os, when Serpophugu cinereu. Torrent Tyrannulet. The streams or rivers that this species requires are not found at the region was much wilder than it is today. Merenberg. Carriker evidently collected at somewhat Mecocerculus minor. Sulphur-bellied Tyrannulet. A higher elevations than Merenbergs’ (16 of relatively rare bird in collections, this species has been the 34 species that he obtained but which recorded from several localities around the head of the Magdalena Valley, at elevations both higher and lower we failed to find are essentially temperate than Merenberg (Meyer de Schauensee 1950). It may zone birds), and his activities were appar- occur here in very small numbers, and possibly was ently concentrated on a ridge west of Santa overlooked. Let&a (a town 12 km NW of Merenberg) Tyrunniscus cinereiceps. Ashy-headed Tyrannulet. and directly connected to the main spine of Status much as for the previous species. Likewise may occur in small numbers at Merenberg. the Central Andes. A last eight species Notiochelidon murinu. Brown-bellied Swallow. Al- (Metallura tyrinnthina, Myiotheretes fu- though common over open and semi-open areas in both migatus, Ochthoeca diadema, Tyranniscus the temperate and piramo zones in Colombia (Meyer nigrocapillus, Cacicus leucorhamphus, de Schauensee 1951, pers. observ.), this swallow al- most certainly would never occur as low as Merenberg. Diglossa caerulescens, Hemispingus atro- Myioborus ornutus. Golden-faced Redstart. Corn- pileus, and Atlapetes !!eucotis) were found mon in temperate forests of Purack National Park, and at similar elevations on the ridge above Is- also seen as low as 2,400 m above Isnos, this redstart nos. This last total could have been raised replaces M. miniutus at higher elevations. An obvious by more prolonged observations. Most of and easily identified bird, we feel certain that it does not range as low as Merenberg. these birds are also primarily temperate Iridosornis rufivertex. Golden-crowned Tanager. zone snecies, and only two (Turanniscus Typically a tempkrate zone bird, and fairly common at nigrocapillus ’ and Diglossa la&ulescens) Puraci, National Park, this tanager certainly would nev- seem at all likely to ever occur at Merenberg. er occur as low as Mere&erg. Anisognuthus Zachrymosus. Lacrimose Mountain- The total thus comes to 174 species. Only Tanager. Another common temperate zone tanager, two other similarly thorough avifaunal stud- 390 ROBERT S. RIDGELY AND STEVEN J. C. GAULIN ies seem to have been reported for single migrants) is dependent on forest, with the areas in the Colombian Andes. Both of these remainder being waterbirds (only two took place on the eastern slope of the West- species, one of them a North American mi- ern Andes in Valle Department, both cen- grant) and birds of open or semi-open areas tering on elevations somewhat lower than (24 species). Of the last, 15 are essentially Merenberg (ca. 2,000 m). Miller (1963) re- tropical zone inhabitants which have been corded 167 species at San Antonio, while S. able to spread upward subsequent to de- Gniadek (unpubl. data) found 176 at Pi- forestation, while 6 occur widely at all chinde (ca. 30 km S of San Antonio). By elevations, and only 3 have been able to comparison, equally intensive studies at lo- spread downslope, only 1 of them at all calities in the humid tropical zone have re- regularly. ported many more species. None has been The human impact on the Merenberg re- reported on from Colombia, but two other gion and its avifauna has also been substan- sites elsewhere in the upper Amazon basin, tial. Extensive areas have been at least par- Limoncocha in eastern and the Ex- tially deforested over the last few decades, plorers’ Inn area near Puerto Maldonado in and the region now supports a rather dense southeastern , appear to be the richest rural population. Many of the larger or rarer localities in the world for bird species. Well birds, especially those dependent on exten- over 450 species have been recorded at Li- sive and continuous forest, have declined or moncocha (Tallman and Tallman 1977, even disappeared altogether. Especially pers. observ.), while the total for the Ex- striking is the absence of some “expected” plorers’ Inn recently surpassed 500 species raptors and large frugivores. Raptors always and is still increasing (T. Parker, pers. seem scarce in the montane forests of South comm.). America (pers. observ.), most species being What accounts for the lower avian diver- relatively secretive and all occurring at low sity found at subtropical elevations such as densities under seemingly optimal condi- Merenbergs?’ Judging from the other two tions. They are also difficult to collect; Car- Colombian studies, Merenberg cannot be rikers’ failure to obtain many comes as no considered atypical. Although subtropical surprise to us (conceivably he may have re- forests are structurally about as complex as frained from shooting them due to their lowland forests, they seem generally to be bulky size, etc.). We do feel that our not hav- less diverse than the latter. This is true of ing recorded raptors locally accurately re- plants (Walter 1971), as well as birds. Ob- flects their current rarity in the region; viously species diversity will, to some seeing them, however, is far easier than col- extent, affect bird species diversity, due to lecting them. Carriker did collect sizeable its effects on the availability of niches, but series of several large frugivores, species this explanation begs the question. Of more such as Ognorhynchus icterotis, general significance may be the fact that, at fuscocinereus, Pyroderus scutatus, and least in Colombia, the humid subtropics, Rupicola peruviana. None of these now oc- where undisturbed, provide but one impor- curs at Merenberg, and it seems certain that tant habitat, the forest. There are no natural their absence can be traced to the activities open areas, relatively little edge or second- of man, both direct (hunting) and indirect ary growth, and almost no open standing (forest clearance, creating patches of forest water. Even where man creates the first and too small and isolated to support these second, these seem to remain under-used. species). However, a few large frugivores To some extent, then, the large avifaunas (e.g., the guan Chamaepetes, the toucan recorded for lowland localities simply re- Andigena nigrirostris, and the two quetzals flect the greater diversity of natural habitat Pharomachrus spp.) have managed to per- there. Lowland areas also appear to have sist. Why these birds have been able to do many more long-distance migratory species so, while the others have not, is an intrigu- (only seven were found at Merenberg). ing question. Comparing places only in terms of their res- Although Merenbergs’ avifauna is some- ident forest avifaunas would probably be what diminished, it remains of unusual in- more meaningful, but in practice would also terest, and the site itself is exceptional in be much more difficult. Slud (1976), for ex- that it provides relatively straightforward ample, did not attempt to distinguish be- access (and for the hardy, even accommo- tween the “forest” and “non-forest” com- dations) to the generally difficult-to-reach ponents of his various locality lists. Of the subtropical forest. We can think of no more Merenberg avifauna, 85% (148 species, in- appropriate way of closing than to commend cluding six of the seven North American the Buch for having had the vision to BIRDS OF FINCA MERENBERG 391 protect their land, and to further hope that Sickle-winged Guan in Colombia. Auk 93:194- the property will be preserved in perpetuity, 195. ideally under the aegis of private conser- KING, W. 1979. Red data book. Vol. 2, Aves. Int. Union Conserv. Nat., Morges, Switzerland. vation organizations such as the recently LEHMANN, V. F. C. 1957. Contribuciones al estudio founded Fundackin Merenberg. de la fauna de Colombia. XII Noved. Columb. 3: 101-156. MEYER DE SCHAUENSEE, R. 1948-1952. The birds of ACKNOWLEDGMENTS the Reuublic of Colombia. Caldasia 5:251-1214. MEYER DE~SCHAUENSEE, R. 1964. The birds of Co- We thank the Buch family for their hospitality at Finca lombia. Livingston Publ. Co., Narbeth, PA. Merenberg and for providing August-December rain- MEYER DE SCHAUENSEE, R. 1966. The species of fall data. Jorge I. Hernandez C. aided us in obtaining birds of . Academy of Natural Sci- the necessary research permits, while Gustav0 Lozano ences, Philadelphia. C. and Roberto Jaramillo M. helped us tremendously MEYER DE SCHAUENSEE, R. 1970. A guide to the birds by identifying much plant material. C. K. Gaulin and of South America. Livingston Publ. Co., Wynne- J. D. Ridgely assisted us in collecting data and in other wood, PA. ways. R. Pasquier at the National Mu- MEYER DE SCHAUENSEE, R., AND W. H. PHELPS, JR. seum and J. Fitzpatrick at the Field Museum in Chi- 1978. A guide to the birds of Venezuela. Princeton cago compiled lists of Moscopin specimens at their Univ. Press, Princeton, NJ. respective institutions, while M. Clench aided us in MILLER, A. H. 1947. The tropical avifauna of the up- compiling such a list at the Carnegie Museum in Pitts- per Magdalena Valley, Colombia. Auk 64:351-381. burgh. S. Hilty contributed useful comments on an ear- MILLER, A. H. 1963. Seasonal activity and ecology of lier draft of this paper, as did two reviewers, E. Eisen- the avifauna of an American equatorial cloud for- mann and J. P. ONeill.’ R. Andrews drew Fig. 1. We est. Univ. Calif. Publ. 2001. 66:1-74. thank all these individuals for their time and expertise. ONEILL,’ J. P., AND T. A. PARKER, III. 1977. Taxon- This study was carried out in the course of larger scale omy and range of Pionus seniloides in Peru. Con- research funded by World Wildlife Fund grant no. US- dor 79:274. 61 (to RSR) and National Science Foundation grant no. RIDGELY, R. S. 1976. A guide to the birds of . BMS75-18145 (to SJCG). Princeton Univ. Press, Princeton, NJ. RIPLEY, S. D. 1977. Rails of the world. D. R. Godine, LITERATURE CITED Boston. ALVAREZ, H. 1975. The social system of the Green SLUD, P. 1976. Geographic and climatic relationships Jay in Colombia. Living Bird 14:544. of avifauna with special reference to comparative BORRERO, J. I. 1952. Algunas aves raras en la Sabana distribution in the neotropics. Smithson. Contrih. de Bogoti. Lozania 1:7-12. 2001. 212: 1-149. BROWN, L., AND D. A~~ADON. 1968. Eagles, and TALLMAN, D. J., AND E. J. TALLMAN. 1977. Adiciones falcons of the World. Country Life Books, Hamlyn y revisiones a la lista de la avifauna de Limonco- Publ. Group, Filtham, Middlesex. cha, Provincia de Napo, Ecuador. Rev. Univ. Ca- CHAPMAN, F. M. 1917. The distribution of bird-life in tolica 5:217-224. Colomhia: a contribution to a biological survey of TODD, W. E. C., AND M. A. CARRIKER, JR. 1922. The South America. Bull. Am. Mus. Nat. Hist. 36:1- birds of the Santa Marta region of Colombia: a 729. study in altitudinal distribution. Ann. Carnegie CHIPLEY, R. M. 1976. The impact of wintering mi- Mus. 14:iii-611. grant Wood Warblers on the resident insectivorous VAURIE, C. 1971. Notes systematiques sur de furna- in a subtropical Colombian oak woods. riides pares des genres Philydor et Xenerpestes, et Living Bird 15:119-141. parallelisme de la forme du bet au type Xenops. FORSHAW, J. M. 1973. Parrots of the World. Lans- LOiseau’ 41:117-126. downe Press, Melbourne, Australia. WALTER, H. 1971. Ecology of tropical and subtropical HAFFER, J. 1974. Avian speciation in South America. vegetation. Transl. by D. Mueller-Dombois. Oliv- Publ. Nuttall Ornithol. Club 14. er and Boyd, Edinburgh. HILTY, S. L. 1977. Chlorospingus flaoovirens redis- WILLIS, E. 0. 1966. Competitive exclusion and birds covered with notes on other Pacific Colombian and at fruiting trees in western Colombia. Auk 83:479- Cauca Valley birds. Auk 94:4449. 480. HILTY, S. L., AND D. SIMON. 1977. Azure-rumped Tanager in Mexico with comparative remarks on RARE, clo Nutional Audubon Society, 950 Third Ave- Gray-and-gold Tanager. Auk 94:605-606. nue, Neu: York, New York 10022. Address of second HOLDRIDGE, L. 1967. Life zone ecology. Tropical Sci- author: Department of Anthropology, University of ence Center, San Jo&, Costa Rica. Pittsburgh, Pittsburgh, Pennsylvania 15260. Accepted JOHNSON, T. B., AND S. L. HILTY. 1976. Notes on the for publication 23 August 1979.