PHAETHORNIS KOEPCKEAE, A NEW OF FROM PER0

JOHN S. WESKE

AND JOHN W. TERBORGH

The Phaethornis comprises a group of dered on each side by a buffy gray malar stripe; sides more than 20 species of predominantly forest- of neck grayish tan; breast light rufescent buff centrally, darker and grayer laterally, and grading dwelling , some of which have uosteriorlv into rich rufescent buff (near 7.5 YR limited ranges and are not well-known. Only 6/8) on abdomen, flanks, and under tail coverts. recently, three new species were named from Remiges and upper primary coverts glossy blackish; Brazil (Grantsau 1968, Ruschi 1972, 1973). upper secondary coverts glossy greenish-bronze. Cen- tral pair of rectrices elongate with slender, 3 mm wide Here we report the discovery of yet another tips that extend 27 mm beyond the tips of adjacent undescribed species (pictured in frontispiece) rectrices; central rectrices dark glossy green basally, in a mountainous area of eastern Peru be- grading through dark grayish subterminally to white at the tips; the four lateral pairs of rectrices gradu- tween Rio Pachitea and Rio Ucayali, and in ated, dark glossy green basally and black subterminally the Maranon Valley of northern Peru. with a broad, buffy rufous (7.5 YR 6/6) tip on each vane. Soft part colors (in life): bill black except orange-red for proximal two-thirds of mandible, iris Phaethornis koepckeae sp. nov. dark brown, feet flesh-color (not blackish as in plate). MEASUREMENTS OF THE HOLOTYPE (mm). KOEPCKE’S -Chord of wing, 58.1; tail (from insertion of the two HOLOTYPE.-American Museum of Natural His- central rectrices to the tip of the longest rectrix), 60.0; tory No. 820872; adult female from the Cerros de1 exposed culmen, 32.6; weight, 5.0 g Sira. 9”3OS.’ 74”47W.’ elevation 860 m, Demo. VARIATIONS AMONG PARATYPES.-No sexual Huanuco, P&6; 2 July ’ 1969; collected by John S. dimorphism is apparent in coloration of the nine Weske and John W. Terborgh; prepared by Weske; specimens at hand. Ventrally the are quite uni- original No. 2015. form, varying only slightly in the intensity of ochra- DIAGNOSIS.-A medium-sized Phacthornis of the ceous color of the abdomen and crissum. Dorsally straight-billed group at one time separated in the there is somewhat more variation, probably age- genus Am&o&s. Nearest to P. philippii (Bourcier) related. In the four birds thought to be adults, the of western Amazonia but slightly larger (see table 1 ), bronzy dorsal area is tinged with rufous only on the with chin and medial throat area white, moustachial rump. In the five birds considered immatures, fine streak whitish, and lateral throat area and sides of rufescent tipping is present on the sides of the crown, neck buffy gray (instead of entire throat, malar area, on the hind neck, and on the lower back; and the and sides of neck rich rufescent buff, shading to rump feathers have broad rufous edges. The tertials oaler buff on chin). It is also somewhat similar of these specimens also have narrow buffy tips that L to P. superciliosus ucayalii, with which it is sympatric, contrast with the blackish color of the rest of the but koepckeae is smaller, with bill shorter and feather. Three of the five show the bill ridges indica- straighter, throat more broadly white, posterior under- tive of immaturity in hummingbirds ( Ortiz-Crespo parts rich rufescent buff ( instead of grayish buff ), and 1972 ) . lateral rectrices tipped with buffy rufous (instead of ETYMOLOGY.-We name this species in memory white ). of our esteemed friend and colleague Dr. Maria DISTRIBUTION.-Recorded at elevations from 690 Koepcke. Before her tragic death in an airplane crash m to 1,130 m on the southwestern slopes of the Cerros in 1971. she had devoted nearlv 20 vears to studvine:, - de1 Sira, in the Provincia de Pachitea, Departamento the birdlife of Peru, making many notable contribu- de Huinuco, east-central Peru. This area lies in the tions. Moreover, by participating in the first two watershed of the Rio Llullapichis, a tributary of the Sira expeditions and by preparing four specimens of Rio Pachitea. Also found in-the Rio Maranon Valley the type series, she was instrumental in the discovery at ca. 506 m elevation near Nazareth and Chiriaco, of the new . Therefore, we feel it is especially Provincia de Bagua, Departamento de Amazonas, fitting to name it in her honor. northern Peru. DESCRIPTION OF THE HOLOTYPE.-Crown REMARKS blackish with a greenish gloss, the feather edges black and giving a scalloped effect; hind neck glossy Field parties from the Louisiana State Uni- greenish-bronze; back glossy bronze (2.5 Y 5/6) (no- versity Museum of Zoology recently encoun- tations refer to colors from Munsell Color Co., 1929- 1942 ): rump is like back but slightly tinged with tered Phaethornis koepckeae in the Maranon rufous; upper tail coverts rufous with two or three Valley in northern Peru. Through the kind- blackish subterminal bars. Lores and auriculars black- ness of John P. ONeill,’ the two LSUMZ speci- ish; short postocular streak white with rufescent mens were available to us as we prepared this tinges; suborbital streak (from base of bill to below auriculars) buffy white; chin and throat white, bor- description. Although they come from a local-

D431 The Condor 79:143-147, 1977 Koepcke’s Hermit (Phaethornis koepckeae), a new species of hummingbird from Per&. Painting by John P. O’Neill. 144 JOHN S. WESKE AND JOHN W. TERBORGH

TABLE 1. Measurements in millimeters of two species of Phaethomis.

P. koepckeae P. philippii

Measurement N Ml2all SD Range N Mean SD Range

Females Wing chord 6 58.3 1.2 57.0-66.1 17 56.4 1.5 53.7-59.9 Tail length 6 61.4 1.1 60.0-62.5 13 59.3 2.9 54.0-64.8 Exposed culmen 6 31.8 1.2 29.5-32.6 17 30.0 0.9 28.0-31.7

Males Wing chord 1 64.6 - - 18 61.2 2.4 57.5-65.9 Tail length 1 64.3 - - 10 62.7 2.8 58.5-68.6 Exposed culmen 2 34.6 - 33.5-34.4 18 32.1 1.6 28.1-34.4

Symbols: N = Sample size, SD = Standard deviation.

ity 600 km northwest of the Sira range, they are known from the Sira area, and two more- are indistinguishable from Sira birds, and we P. hispidus and P. philippii-probably occur have no hesitation in including them among at its eastern base along the Rio Ucayali, an the paratypes. area we did not visit. Consideration of the elevational ranges and relative abundance HABITAT AND ECOLOGY (fig. 1) of the Phaethornis we encountered allows conjecture about their competitive rela- The Cerros de1 Sira comprise a small, isolated tionships. Phaethornis superciliosus ucayalii range of mountains that rises abruptly from is abundant in lowland forest and ranges up- the Amazonian plain 60 km to the east of the ward through hill forest approximately to the main Andean Cordillera. The elevation of lower limit of cloud forest. In the lowlands the highest peak is about 2,400 m. The area, it coexists with P. longuemareus atrimentalis, which has been described in detail by Ter- a species weighing only half as much. Phae- borgh and Weske ( 1975), includes the sole thornis longuemareus is probably more com- known Peruvian locality for the Horned Cu- mon than figure 1 suggests, because its tiny rassow (Pauxi unicornis; see Weske and Ter- size causes it to be ineffectively sampled by borgh 1971). mist-nets. Phaethornis superciliosus and P. The habitat of Phaethornis koepckeae in the koepckeae occur together in the foothills. The Sira is the understory of humid lower mon- latter weighs about 16% less than the former, tane forest (“hill forest”). Although the spe- and its straighter bill averages 13% shorter. cies was found in small numbers in lowland- These differences in size and morphology type forest at 690 m and once in mossy cloud may be correlated with different feeding forest at 1,130 m, it was most common in the niches, but we have no behavioral information zone just below the lower limit of cloud that confirms their segregation in this manner. forest (fig. 1). At this elevation the forest David L. Pearson (in litt.) found that in understory is quite dense and contains large Ecuador, the “Ametrornis” hummingbird populations of Zingiberaceae (, Reneal- Phaethornis bourcieri fed higher in the forest mia), Marantaceae (Calathea), and Musa- and more frequently on insects than did P. ceae ()-plants that are known to superciliosus and P. longuemareus at the same serve as food sources for hermits. site. According to Theodore A. Parker III (pers. Single individuals of P. superciliosus and comm.), the habitat of P. koepckeae in north- P. koepckeae were captured at 1,130 m, a little ern Peru is very‘ moist upper tropical for- above the lower limit of cloud forest in the est”-similar, apparently, to its habitat in the Sira. At this camp the most frequent Phae- Sira. The collecting elevation on the label of thornis was P. guy apicalis, which weighs one of the Maranon Valley specimens is 1,700 virtually the same as P. koepckeae but has a ft (520 m), not inconsistent with the Sira long curved bill like P. superciliosus. The data although a little lower, but for the other bird suggest that these three congeners, all quite it was listed as 900 ft (270 m). However, similar in size, cannot coexist effectively. Parker assures us that this locality (km 381.4), which he has visited, is actually considerably Competition from P. superciliosus and P. higher and that the two Maranon Valley speci- koepckeae may combine to exclude P. guy mens came from roughly the same elevation, from hill forest elevations. In the Apurimac about 1,700 ft. Valley of Peru, where P. koepckeae is absent, Four species of Phaethornis hummingbirds P. guy ranges from 1,700 m down to the lower NEW PERUVIAN HUMMINGBIRD 145 limit of hill forest at 685 m, overlapping broadly with P. superciliosus between 685 m 004. and 930 m ( Weske 1972). On Trinidad, where it is the only large Phaethornis, P. guy occurs commonly at sea level as well as higher (J.W. 003.

T., pers. observ.). $ In the Sira, peak numbers of P. koepckeae 5 gooz- occurred at or near 860 m, where among 70 bird species captured in mist-nets it ranked s (with the hummingbird Thalurania furcata) 0.01 as the fifth most common species. The four most abundant species in this sample, in des- t OOOL ’ I cending order, were the manakin Pipra pipra, 200 400 600 800 1000 1200 140 ELEVATION (METERS) the flycatcher Mionectes striaticollis, the woodcreeper Glyphorhynchus spirurus, and FIGURE 1. Relative abundance of Phaethornis the manakin Pipra chloromeros. hummingbirds at various elevations in the Cerros de1 Sira. The proportional occurrence of each species in mist-netted samples is plotted along the ordinate. BREEDING AND MOLT The Sira specimens of P. koepckeae were all taken in July, a dry season month, while the widespread species ranging through the west- Maranon Valley ones are from October and ern Amazonian lowlands (fig. 2). Over this December, the early part of the wet season. vast range it shows no appreciable geographic Gonadal examination revealed that none of variation and was considered monotypic by these birds was in breeding condition. How- Zimmer (1950). Likewise, the two known ever, at least a fraction of the population populations of P. koepckeae are undifferen- breeds in July, for Jane Brockmann found tiated. David L. Pearson (in litt. ), who has and photographed a nest with two young on observed philippii near Pucallpa, on the Rio 23 July 1971 at 690 m in the Sira. It was a Ucayali, Peru (elev. 140 m), reports that it is small conical structure located 1.2 m above a bird of mature forest, as is koepckeae. But the ground and attached to the underside of a unlike the latter, which is an inhabitant of hill cyclanth frond near the tip (illustrated in country, philippii is strictly a denizen of the frontispiece). The December specimen is lowlands, not recorded anywhere at elevations molting at least its flight feathers. The outer greater than about 200 m. Its presence near primaries are sheathed basally as are several Pucallpa, 100 km north of the Sira, and at rectrices. The long, central rectrices are about Lagarto, 150 km southeast of the Sira, implies half-grown. continuous distribution along the Rio Ucayali SYSTEMATIC RELATIONSHIPS valley. Since koepckeae can be expected to live throughout the 600 to 1,100 m zone on the Besides P. koepckeae, the “Ametrornis” group slopes of the Sira, the ranges of the two spe- includes P. bourcieri, a smaller, gray and green species found in the lowlands of north- cies probably approach one another within ern South America to northeastern Peru, and several kilometers where the Ucayali borders P. philippii. On morphological grounds alone, the Siras’ eastern side. Sympatry would be one might be inclined to consider koepckeae most unlikely as neither would be expected at and philippii conspecific. They do not differ elevations between 200 and at least 500 m. As greatly in measurements (table l), and they is true in numerous other genera of this region are remarkably similar in bill shape and in (Terborgh 1971, Terborgh and Weske 1975)) plumage pattern and coloration except for the P. koepckeae and P. philippii are two closely prominent differences on the face and an- allied but distinct species which have evolved terior underparts. However, Phaethornis is a different elevational and habitat requirements, quite uniform genus in which certain other recognized species show relatively little mor- SPECIMENS EXAMINED phological differentiation from some conge- Phaethornis koepckeae. PERU. Depto. Amazonas: ners. Moreover, there are compelling 20 km SW Chiriaco, ca. 1,700 ft, 15 Dec. 1974, 1 $ zoogeographic and ecological grounds for con- im. (LSUMZ); km 381.4 on Corral Quemado-Naz- areth Highway, ca. 900 ft, 8 Oct. 1973, 1 9 im. sidering the two forms to be distinct species. (LSUMZ). Depto. Huanuco: Cerros de1 Sira, 650 m Phaethornis philippii is an uncommon but l= 690 m, adjusted], 24-26 July 1971, 1 $ im., 3 146 JOHN S. WESKE AND JOHN W. TERBORGH

FIGURE 2. Distribution of Phaethornis koepckeue (squares ) and P. philippii (circles). Localities are based on Gyldenstolpe (1945, 1951) as well as on specimens examined in this study.

9 o ad.; Cerros de1 Sira, 860 m, 24 July 1969, 1 LITERATURE CITED 0 ad., 1 0 im., 1 unsexed im. (AMNH). GRANTSAW, R. 1968. Una nova especie de Phae- Phaethornis philippii. PERU. Depto. Loreto: Santa thornis ( Aves, Trochilidae). Pap. Avulsos Zool. Cecilia on Rio Maniti, 3 $ $ (FM); 40 mi. E Iquitos, 22( 7) :57-59. 3 $ $, 2 0 9 (FM); Rio Pichana, 1 8, 2 99 GYLDENSTOLPE, N. 1945. The bird fauna of the (FM); Yarinacocha, 1 $ (LSUMZ), 1 0 (FM); 59 Rio Jurua in western Brazil. K. Sven. Veten- km W Pucallpa, 4 $ $ , 4 9 9, 1 unsexed ( AMNH). skapsakad. Handl. Ser. 3,22( 3) : 1-338. Depto. Madre de Dios: 105 km SW Puerto Mal- GYLDENSTOLPE, N. 1951. The ornithology of the donado, 1 unsexed (LSUMZ). BRAZIL. Est. Ama- Rio Purus region in western Brazil. Arkiv. Zoo]. zonas: Nova Olinda on Rio Pm&, 1 $ (ANSP). Ser. 2, 2( 1):1-320. Also the AMNH series from Peni and Brazil listed by MUNSELL COLOR Co., INC. 1929-1942. Munsell Zimmer (1950), including the type. book of color. Baltimore, Maryland, pub]. by Phaethornis superciliosus ucayalii. The AMNH series the author. listed sub nom. P. maluris ucayali by Zimmer ( 1950). ORTIZ-CRESPO, F. I. 1972. A new method to sep- arate immature and adult hummingbirds. Auk 89:851-857. ACKNOWLEDGMENTS RUSCHI, A, 1972. Una nova especie de beija-flor We thank the following for allowing examination of do E. E. Santo, Phaethornis margarettae. Bol. collections in their care or for loan of specimens: Mus. Biol. Mello Leitgo, Zoo]. Ser. 35:1-5. J. P. ONeill’ and G. H. Lowery, Jr., of the Louisiana RUSCHI, A. 1973. Una nova especie de beija-flor State University Museum of Zoology (LSUMZ). W. do E. E. Santo, Phaethods nigrirostris. Bol. E. Lanyon of the American Museum of Natural Mus. Biol. Mello LeitPo, Zoo]. Ser. 36:l3. History (AMNH); E. R. Blake of the Field Museum TERBORGH, J. 1971. Distribution on environmental of Natural History (FM); and F. B. Gill of the gradients: Theory and a preliminary interpreta- Academy of Natural Sciences of Philadelphia (AN- tion of distributional patterns in the avifauna of SP). David L. Pearson, Theodore A. Parker III, the Cordillera Vilcabamba, Peru. Ecology 52: Jane Brockmann, Robert W. Dickerman, and Mary 23-40. H. Clench supplied helpful information on field ob- TERBORGH, J., AND J. S. WESKE. 1975. The role servations or specimens. E. Eisenmann, R. C. Banks, of competition in the distribution of Andean J. W. Fitzpatrick, J, P. ONeill,’ and J. T. Marshall, birds. Ecology 56:562-576. Jr., critically read the manuscript and made useful suggestions for its improvement. Field work in the WESKE, J. S. 1972. The distribution of the avi- Cerros de1 Sira was supported by the National fauna in the Apurimac Valley of Peru with Science Foundation ( GB-20170). respect to environmental gradients, habitat, and NEW PERUVIAN HUMMINGBIRD 147

related species. Ph.D. diss. Univ. Oklahoma, National Fish and Wildlife Laboratory, U.S. Fish Norman. and Wildlife Service, National Museum of Natural WESKE, J. S., AND J. W. TERBORGH. 1971. A new History, Washington, D.C. 20560, and Depatiment of curassow of the genus Pauxi from of Biology, Princeton University, Princeton, New Peru. Auk 88:233-238. Jersey 08540. Accepted for publication 13 Novem- ZIMMER, J. T. 1950. Studies of Peruvian birds. ber 1976. No. 55. Amer. Mus. Novit., No. 1449.