The Auk 116(4):882-890, 1999

A NEW SPECIESOF (FORMICARIIDAE: ) FROM THE SOUTHERN ECUADORIAN ANDES

NIELS KRABBE,TM D. J. AGRO,3 N.H. RICE,4'7 M. JACOME,2 L. NAVARRETE,5 AND E SORNOZA M? •ZoologicalMuseum, University of Copenhagen,Universitetsparken 15, DK-2100,Denmark; 2MuseoEcuatoriano de CienciasNaturales, Casilla 17-7-8976, Quito, ; 3Bird Studies Canada, P.O. Box 160, Port Rowan, Ontario NOE 1MO, Canada; 4Departmentof Systematicsand Ecology and Museum of NaturalHistory, University of Kansas, Lawrence,Kansas 66045, USA; and sCooperativaSanta Marta 1, JuanLeon Mera, SantaDomingo de losColorados, Ecuador

ABSTRACT.--Wedescribe a strikingnew speciesof Grailariaantpitta from wet, uppersub- tropicalforest in the upperRio Chinchipedrainage, provincia Zamora-Chinchipe, Ecuador Noteson its naturalhistory and molecularsystematics are presentedalong with spectro- gramsof its voice.The speciesis readilydiagnosed by its largesize, unique white facial markings,and voice.Initial resultsfrom DNA sequenceanalyses place the new in a well-supportedclade that includesGrailaria nuchalis (which is syntopicwith the new spe- cies),G. hypoleuca,G. watkinsi,and G. ruficapilla.Received 23 September1998, accepted 22 April 1999.

RESUMEN.--Unasorprendente nueva especie del g6neroGrailaria (Aves: Formicariidae) es descritade un bosquehfimedo de la zona subtropicalalta del drenajedel Rio Chinchipe, provinciaZamora-Chinchipe, Ecuador Se presentan notas acerca de su historianatural y sistemfiticamolecular, en conjuntocon espectrogramas de susvocalizaciones. La nuevaes- peciese diferencia claramente por su grantamafio, sus marcas faciales blancas, y supeculiar voz. Anfilisisiniciales de sequenciasdel ADN colocana la nuevaespecie en un cladoque incluyea Grailarianuchalis (que es sint6picacon la nuevaespecie), G. hypoleuca,G. watkinsi, y G. ruficapilla.

FIELDWORK IN SOUTHAMERICA during the speciesof antpitta with strikingly distinct past25 yearshas resultedin the discoveryof plumagecharacteristics was discoveredon 20 many new to science(Mayr and Vuilleu- November 1997 by R. S. Ridgely, J. and R. mier 1983, Vuilleumier and Mayr 1987). New Moore, L. Navarrete, and M. Rivadeneira dur- generahave been describedfor somespecies ing a trip to recordvocalizations of Ecuadorian (e.g. Xenoglauxloweryi, O'Neill and Graves birds. Ridgely,the Moores,and Navarretere- 1977;Acrobatornis fonsecai, Pacheco et al. 1996; turned on 21 November 1997 to record and Nephelornisoneilli, Lowery and Tallman 1976), photographthe , and M. Jicome,Navarre- and the known rangesof other generahave te, X. Mufioz, and F. Sornoza M. visited the site been greatly extendedby other new species on 27 to 29 December 1997 and collected the (e.g.Nannopsittaca dachilleae, O'Neill et al. 1991; first specimen.A team from the Academyof Clytoctantesatrogularis, Lanyon et al. 1990). Natural Sciencesof Philadelphia(ANSP), the Mostdiscoveries, however, have been of species Museo Ecuatoriano de Ciencias Naturales with obviousrelatives and which fit logically (MECN), the ZoologicalMuseum of the Uni- into broaderinterpretations of theirrespective versityof Copenhagen(ZMUC), and others(D. group'shistorical biogeography. J. Agro, A.D. Brewer,N. Krabbe,L. Navarrete, Therefore,it cameas a surprisethat a new R. S. Ridgely,N. Simpson,E SornozaM., and D. L. Wechsler)retumed to the original site be- 6Address correspondence to this authorat Museo tween 13 and 22 January1998 to assesspopu- Ecuatoriano de Ciencias Naturales, Casilla 17-7- lation size and habitatrequirements and to ob- 8976,Quito, Ecuador.E-mail: [email protected] tain further specimensand photographsof the 7Present address: Academy of Natural Sciencesof bird. Sornoza and Jicome obtained an addi- Philadelphia,1900 Benjamin Franklin Parkway, Phil- tional specimenduring a visit to nearby Cerro adelphia,Pennsylvania 19103, USA. Toledo from 16 to 31 March 1998.

882 October1999] NewAntpitta from Ecuador 883

FIG.1. Headof Grailariaridgelyi showing black and white pattern. Inset is magnificationof cheekfeather. Drawing by S. Trammell.

Five specimensof the new bird were collect- arablefrom known antpittasby white cheek ed and comparedwith specimensof otherfor- patch of rigid loose-barbedfeathers with micariids.They clearlyrepresent a new taxon glossyshafts extending from the subocular and that we proposeto name: posteriorpart of the lower loresto the baseof the bill (seeFig. 1). Thewhite cheekpatch con- Grailaria ridgelyi sp. nov. trastssharply with theblack cap, bill malarre- JocotocoAntpitta gion, and anteriorauricular feathers. The new speciesdiffers from all other Grailariain lack- Holotype.--MECNNo. 7199,adult male (skull ing true rictalbristles. The throatfeathers are ossified,no bursa of Fabricius),collected 28 De- white, stiff, and loose-barbed, as in the Ochre- cember1997 by E SornozaM. (FS 1922)in Que- stripedAntpitta (G. dignissima),but are denser, bradaHonda, provincia Zamora-Chinchipe, Ec- shorter,and with more recurvedtips, as in uador (04ø29'S, 79ø08'W; elevation 2,520 m). Chestnut-napedAntpitta (G. nuchalis).The Vocalrecordings are archived at theLibrary of gray of the underpartsis interspersedwith Natural Sounds(LNS 79055).A blood sample white in a pattern resemblingpale-bellied (FS-1-28.12.97)is depositedat the Institutefor membersof the G. hypoleucaallospecies group. PopulationEcology, Copenhagen University The bill is shapedmost like that of G. nuchalis (IPE); tissue samplesare deposited as ANSP and the Pale-billedAntpitta (G. carrikeri) but is 8701. proportionatelydeeper at the baseand with a Diagnosis.--Alarge antpitta (body mass150 straighterculmen. Tarsi and toesproportion- to 200 g) assignableto the genusGrailaria. Sep- atelylarger than in otherantpittas and aslarge 884 KRABBEET AL. [Auk, Vol. 116 as in Giant Antpitta (G. gigantea).Tarsal scutes brown, and the tarsi and toes are blue-gray. few and distinctlyridged with toesand claws The stomach contained a few insect remains, shortand sturdy,resembling G. nuchalis. and someof the remiges,rectrices, and body Descriptionof holotype.--Crown,a 1 to 2-mm featherswere being replaced. wide featheredocular ring, broadmalars, and Measurementsof holotype.--Unflattenedwing anteriorauriculars connecting with malarsJet chord130 mm, flattenedwing chord138 mm, Black (capitalized color names and numbers tail 59.6 mm, bill from anterioredge of nostril follow Smithe1975). A large and distinctfan- 19.3 mm, bill from skull 33.5 mm, gonys18.8 shaped patch of rigid, loose-barbedwhite mm, tarsus67.1 mm, body mass176 g (light featherswith glossyshafts in theloral and sub- fat), testes12 x 3 min. ocularregion (seeFig. 1). Thesefeathers are Designationof paratypes.--Adultmale: ANSP longerand denserthan in congenersand cover 187810(skin); Natural History Museumat Uni- the base of the bill and most of the malar re- versityof Kansas(KU) 89207(trunk alcoholic gion. Rictal bristleslacking, but a few of the specimen);ANSP 8706 (tissue);KU 2214 (tissue white loral feathershave elongated bare shafts,' subsampleand genomicextract); LNS 79054 formingbristle-like extensions. Poorly defined (voice). Adult male: MECN 7284 (skin); IPE postocularstreak, posterior part of auriculars, FS1-31.3.98(blood). Adult female:MECN 7198 andsides of neckLight Neutral Gray (85). Nape (skin); ANSP 8704 (tissue). Adult female: black but suffused with color of back, which is ANSP 187811(skin with all bonesexcept those closestto BrownishOlive (129),but moregold- of right tarsometatarsusand foot removed;i.e. en.Upper mantle with blackwash that decreas- a "schmoo");KU 89206(skeleton); ANSP 8731 es posteriorly.All back featherswith blackish (tissue);KU 2215 (tissuesubsample and geno- shafts.Tail, composedof 12 rectrices,between mic extract);IPE NK1-21.1.98 (blood). Raw Umber (223)and Mars Brown(223A). The MECN 7284 was taken on the east slope of three innermost secondaries and outer webs of CerroToledo (2,680 m) 6 km northof the type the nineother secondaries slightly lighter than locality,just within PodocarpusNational Park, Chestnut(32). The distalhalf of the outerwebs on 31 March 1998.The remainingthree para- of the 10 primaries are Ground Cinnamon typeswere taken near the type localityin Jan- (239),the innerwebs of the remigesare black- uary 1998 at 2,550 m. ish, and the underwing primary covertsare Variationin typeseries.--Males average larger mostlyblack. Upperwing coverts Brownish Ol- than females,although the sexesoverlap in ive with 2-mm wide blackish shaft streaks and measurementsof mostcharacters (Table 1). The Chestnutedges and tips thatare bordered by a belly is white in two malesand white lightly 1-mm wide blackish band. Alula Brownish Ol- suffusedwith LightNeutral Gray in thetwo fe- ive like back,outer webs nearly uniform. Breast males and the third male (MECN 7284). The White with light suffusionof Light Neutral amount of buff on the crissum is variable and Gray. Throat white, featherswith fairly stiff, is strongestin two of the males,fainter in the glossy,elongated and recurvedshafts that on other male (ANSP 187810) and one female some chin feathersare faintly dark-tipped. (ANSP 187811),and absentin the otherfemale. Feathersof upper breastwith faint, blackish, The faintbars on theupper breast are most dis- about l-ram wide terminal suffusions. Crissum tinctiveon the holotypeand MECN 7284.The and extreme lower belly washed with Pale amount of black suffusion on the mantle is least PinkishBuff (121D). Sidesof breastBrownish extensivein the holotype.Noticeable variation Olive.Sides and flanksLight NeutralGray suf- existsin the intensityof the grayon thebreast. fusedwith BrownishOlive. ThighsBrownish Amongthe paratypesthe bill is jet black,with Olive admixedwith Light Neutral Gray and the tip of the mandiblegrayish in one female heavilywashed with black.Undertail coverts (ANSP 187811).Other soft-part colorswere and underwing covertsBrownish Olive, with (ANSP 187810 [male] and MECN 7198 [fe- fineblack bars. A singlecompletely black feath- male]):irides Kingfisher Rufous (240), tarsi and er is on the left shoulder,a nearlyblack one on toes Pratt's Payne'sGray (88) with blackish the right thigh, and most of the olive-brown joints and edgesof scutes,nails Dark Neutral feathershave faint blackish tips. In life, thebill Gray (83); ANSP 187811 (female): mouth is jet black, the irides are crimsonreddish creamywhite tinged pink, irides Mahogany October1999] NewAntpitta from Ecuador 885

TABLE1. Measurementsof Grailariaridgelyi sp. nov. Values are œ + SD of threemales and two females (range in parentheses).

Variable Males Females Body mass(g) 192 + 14.3 (176 to 204) 167 +_21.2 (152 to 182) Exposedculmen (mm) 28.2 +_1.0 (27.1 to 29.0) 27.8 + 1.1 (27.0 to 28.5) Bill from skull (mm) 34.8 _+1.63 (33.5 to 36.6) 33.0 -+ 0.14 (32.9 to 33.1) Bill from nostril (mm) 19.5 + 0.85 (18.9 to 20.5) 18.4 +_0.35 (18.1 to 18.6) Bill width at base(mm) 16.1 + 1.01(15 to 17) 16.0+_ 0.28 (15.8to 16.2) Gonys(mm) 19.8 _+1.27 (18.8to 21.2) 18.9 + 0.14 (18.8to 19.0) Unflattenedwing chord(mm) 131_+ 3.61 (128 to 135) 126-+ 2.83 (124 to 128) Flattenedwing chord(mm) 138 +_2.52 (135to 140) 132 + 1.41(131 to 133) Tail (mm) 58 + 2.43 (55 to 66) 54 +_4.81 (50 to 57) Tarsus(mm) 71.1 _+3.59 (67.1 to 74.1) 70.4 -+ 4.24 (67.4 to 73.4) Middle toe (mm) 47.1 _+1.91 (46.0 to 49.3) 45.4 (n = 1) Hind claw (mm) 13.9 + 0.25 (13.6 to 14.1) 13.6 _+0.57 (13.2 to 14.0)

Red (132B), tarsi Plumbeous Gray (87-88); transcriptionof the Spanish"Jocotoco" (pro- MECN 7284 (male): irides maroon, feet bluish nounced"hocot6co"), a localonomatopoeia for gray. the antpitta. MECN 7284: testes 13 X 5 mm, seminal ves- iclesenlarged; ANSP 187810:testes 5 X 2 mm; REMARKS MECN 7198: ovary 11 X 6 mm, largest ovum 1.8 mm; ANSP 187811:ovary 15 X 3 mm, larg- Population.--Thatsuch a largeand distinctive est ovum 2 mm, oviduct convoluted. speciesremained undiscovered until 1997sug- Distribution.--Known from a small area of geststhat G. ridgelyihas a very limited geo- very wet forestin the upper Rio Chinchipe graphicdistribution and/or specifichabitat re- drainage,provincia Zamora-Chinchipe, south- quirements.At the type locality,we estimated ern Ecuador, where it has been recorded at el- six territoriesof G. ridgelyiper squarekm and evationsranging from 2,300 to 2,680 m. notedthat seeminglysimilar continued Comparativematerial.--We examined all to the southeastand southwest.Accordingly, knownspecies of Grailariawith the exceptionof we speculatethat it occursthroughout these the TachiraAntpitta (G. chthonia),for whicha forests,ranging east and southeastto Cordi- suitabledescription is available(specimens in llera de Tzunantza and the southernpart of MECN, ANSP,and LouisianaState University Cordillera del Condor, and southwest to the Museum of Natural Science[LSUMNS]). Rio Isimanchi.We considerit possiblethat, de- Etymology.--Weare pleased to namethis spe- spiteprevious surveys that failedto detectthe ciesin honorof RobertS. Ridgely.During his species(Parker et al. 1985,Rasmussen and Rah- field work in South America over the last 35 bek 1994),G. ridgelyialso ranges farther south- years,and duringhis manyyears at the Acad- westto CerroChinguela, Peru, and throughout emy of Natural Sciencesof Philadelphia,Bob the PodocarpusNational Park north to Rio Za- hasgathered a vastknowledge of Neotropical mora.Its rangemight even exceed these limits, birds. His combination of field and museum ex- and we encouragesearches for it as far north perienceand thorough acquaintance with Neo- and south as Volc•n Sumaco in northern Ec- tropical ornithologicalliterature is outstand- uador and the centralAndes of Peru,respec- ing. He hasshared his knowledgewith a wide tively. audience and stimulated an interest in Neo- Habitat.--Thefive specimenswere collected tropicalbirds through his booksA Guideto the in wet montaneforest with two or morespecies Birds of Panama(Ridgely and Gwynne 1976) of bambooon steepslopes in the upper sub- and TheBirds of SouthAmerica (Ridgely and Tu- tropicalzone. The highestelevation at which dor 1989,1994). It is fitting that the discovery the birds were found (2,680m) coincideswith of thisastounding new antpitta comes as work the upperlimit of Cecropiatrees, which are a with his coauthorPaul J. Greenfield on yet an- much-usedindicator of the upper limit of the other landmark book, The Birdsof Ecuador, subtropicalzone (Ulloa and Joergensen1993). nearscompletion. The Englishname is a direct Tree height rarely exceeded20 m on steep 886 KRABBEET AL. [Auk,Vol. 116 slopes,with most trees about10 m tall. Most threwtheir heads back in typicalantpitta fash- trunks, stems, and branches were covered in ion. Theyusually responded in closepairs, as thicklayers of dripping-wetmoss, lichens, and do severalother members of the family,notably liverworts. During our 12 field days we fre- someGrallaricula and Grailarianuchalis (Fjelds• quentlyexperienced rains and heavyfog. and Krabbe 1990). Vocalizations.--Thesong of G. ridgelyi(Fig. Diet.--The five stomachs contained "a few 2A), whichwas only observedgiven by males, insect remains," "worms and ," "ants," consistsof a seriesof 6 to 10 or more notes (400 " parts and larvae,.... re- to 650 Hz) separatedby intervalsof 1 to 2 s, dis- mains," and "millipede and insectremains." tinctly lowerpitched than songsof its closest All stomachcontents have been depositedat relatives(Fig. 2D). The qualityof the notesis ANSP for future identificationof specificprey reminiscentof thoseof Pteroptochostapaculos items. and the Rufous-bandedOwl (Strix albitarsis), Breeding.--Nospecimen had greatly en- and at a distancethe notesmay be confused largedgonads, all had light fat reserves,and all with a barkingdog. Following playback, notes but one,a heavilyworn January female, were in weregiven for aslong as 90 s;the pitchand am- generalmolt (presumablyprebasic); this sug- plitude (volume)rose slowly for more than 1 geststhat breedingoccurs at leastin October min, with notesdelivered every 0.6 s;the pitch and November, which is consistent with the then leveledand the pacebecame slower and breeding cycle of congenersin this region more dog-like. (pers.obs., M. B. Robbinspers. comm.).How- The call (Fig. 2B), givenby both sexes,is a ever,despite being in molt and havingfairly softer, two-noted "ho-co." Notes are about 575 smalltestes, the Marchspecimen had well-de- and 475Hz, with the secondnote occasionally veloped seminalvesicles. higherpitched. We heardthe call givensingly, or repeatedlyevery 4 to 10 s, with intervals SYSTEMATICS shortestafter playback. The alarm (Fig. 2C), given after repeated The taxonomicaffinities of the new species playback,is similarto the call but the lastpart werestudied through phylogenetic analysis of is a churred "h6-crrr" or "ho-c6-crrr." mitochondrialDNA by N.H. Rice. Songsand callsgenerally were givenat dawn Taxa examined.--Elevenspecies of Grailaria and duskbut alsoat othertimes of day during antpittaswere studied,including two individ- rain and in misty weather Birdsvocalized con- ualsof thenew species (Table 2). Previouswork siderably more in November and December (N. Riceunpubl. data) suggestedthat Grallari- than in January,when natural song was heard cula, Hylopezus,and Myrmotheraform a sister on only a few occasions.In late March, we lineageto Grailariaand that togetherthese four heardsongs only afterplaybacks. genera form a well-supported monophyletic Behavior.--Thespecies has only been ob- group. Accordingly, single representatives served after respondingto playbackof songs from Grallaricula,Hylopezus, and Myrmothera and calls.Most individuals (four pairs and one were chosenas outgroupsfor characterpolar- solitary bird) approachedconsiderably less ization(Table 2). Freshlyfrozen or ethanol-pre- cautiouslythan most other species of Grailaria, servedtissues (liver, heart, and muscle)with and they were often heard moving through associatedvoucher specimens were obtained vegetationor occasionallyon the forestfloor from LSUMNS and ANSP. beforethey were heard singing. The birds were Phylogeneticresults.--The cytochrome-b and relatively aggressive,sometimes coming to ND2 matricesproduced congruenttree topol- within 4 or 5 m of the observerThey moved ac- ogieswith comparablesupport regardless of tivelyfrom branch to branch,or ran alonghor- weightingscheme. For brevity,therefore, only izontalbranches 2 to 4 m abovethe ground (less thetopology resulting from the combined anal- oftenon the forestfloor), periodically stopping ysis is discussedin detail. to sing. Occasionally,excited individuals rap- UsingGrallaricula, Hylopezus, and Myrmothera idly bobbedtheir headsto below the level of as outgroups,and with equal weightingas- their perch.Singing birds pumpedtheir tails signed to transitions and transversions,a sin- down in synchronywith each "hoot" and gle most-parsimonioustree resulted (TL = October1999] NewAntpitta from Ecuador 887

A

I I

B

0 i I

c

1-

i I

D

i i 10 20

Time ($) FIG.2. (A) Songof G. ridgelyi(LNS 88163;recorded by L. Navarrete).(B) Male call of G. ridgelyi(recorded by D. Agra). (C) Excitedmale call of G. ridgelyiafter repeatedplayback of song(recorded by D. Agra). (D) Left to right, songsof G. ridgelyi(see Fig. 2A), G. carrikeri(La Libertad,Peru; recorded by T. A. Parker),G. nuchalis(Napa, Ecuador;recorded by N. Krabbe),G. watkinsi,and G. ruficapilla(Laja, Ecuador;recorded by N. Krabbe),respectively.

1,261, CI = 0.548, HI = 0.452, RI = 0.518; Fig. (1) large antpittasof the subgenusGrailaria 3). This hypothesissupports monophyly of (Loweryand O'Neill 1969),here represented by Grailaria.Within Grailaria,it recognizestwo G.squamigera, G.varia, and G.guatimalensis; and well-supportedand strongly resolvedclades: (2) antpittasreferred to the subgeneraTham- 888 KRABBEET AL. [Auk, Vol. 116

TABLE2. Taxaand tissue samples used in thisstudy, with GenBankaccession numbers of thederived DNA sequences.

GenBank number Taxon Tissue no. ND2 Cytochromeb Grailariasquamigera LSUMNS 6254 AF127203 AF127188 G. varia LSUMNS 7528 AF127204 AF127189 G.guatimalensis ANSP 5173 AF127205 AF127194 G. rufula LSUMNS1218 AF127206 AF127190 G. blakei LSUMNS 5620 AF127207 AF127191 G. ruficapilla ANSP 4906 AF127208 AF127192 G. watkinsi ANSP 2906 AF127209 AF127193 G. nuchalis ANSP 4905 AF127212 AF127197 G. ridgelyi1 ANSP 8706 AF127210 AF127195 G. ridgelyi2 ANSP 8731 AF127211 AF127196 G. hypoleuca ANSP 4476 AF127213 AF127198 G. dignissima ANSP 3229 AF127214 AF127199 Hylopezusfulviventris ANSP 3744 AF127216 AF127202 Grallariculalineifrons ANSP 4039 AF127215 AF127200 Myrmotheracampanisona LSUMNS 7563 AF127217 AF127201

nocharis,Hypsibemon, and Oropezusby Lowery Hypothesisinterpretation and implications.- and O'Neill (1969),although this division is not The new speciesis strongly supported as a supported.The cladehas the G. rufulacomplex member of the Grailariaclade (97% bootstrap (G. rufulaand G. blakei)as its basallineage, and value) and, within the limits of our taxon sam- amongthe other speciesan unexpectedsister pling,as the sisterto G.nuchalis (82% bootstrap speciesrelationship between G. hypoleucaand support).From this relationship,we infer that G.dignissima. Grailaria ruficapilla and G.watkinsi G. carrikeri, for which no biochemical material are sistertaxa and surprisingly,they form the is available,but which Schulenbergand Wil- sisterlineage to a clade containingG. ridgelyi liams (1982) suggestedis a relative of G. nu- and G. muchalis. chalis, also is a member of the clade that con-

Gralla#asquamigera

0 1O07 • G.vat•a G. guatimalensis 97 G. rufu/a 97 125 l__ G.b/akei 1O0 G.ru•capilla 80 2 1 • G. n'dgelyi 82 G. ridgelyi 99 8 3 4 3 G.nucha/is

796 5 (• G. dignissirnahypo/euca

<50 Hylopezusfulviventris 0 0 -- Grallat•culalineifrons Myrrnotheracarnpanisona F•G.3. Cladogramrepresenting the singlemost-parsimonious tree based on combinedmolecular data. Numbersabove each node are bootstrap values for eachclade, and numbers below each node are decay in- dices(left) and numberof unreversedsynapomorphies (right) for eachclade. October1999] NewAntpitta frorn Ecuador 889 tains G. ridgelyiand G. nuchalis.Putative mor- portfor thelaboratory portion of thisstudy was pro- phologicalsynapomorphies supporting this re- videdby a FrankM. Chapmangrant from the Amer- lationship(carrikeri + nuchalis+ ridgelyi)are ican Museumof Natural History to NHR. N. Simp- that all haveunpatterned breasts, smoky gray son generouslysponsored the establishmentof flanks,and a deep,robust bill. Consistentwith "Fundaci6nJocotoco" and the purchaseof the type locality. this is their biogeography:all three have an eastern Andean distribution concentrated in northern Peru, Ecuador, and . Inter- LITERATURE CITED estingly,G. carrikeri and G. ridgelyi have distinct FJELDS,g,,J.,AND N. KRABBE.1990. Birds of the High blackcaps and reddish irides. Andes. ZoologicalMuseum, University of Co- penhagen,Copenhagen, Denmark. CONSERVATION LANYON, S. M., D. E STOTZ,AND D. E. WILLARD. 1990. Clytoctantesatrogularis, a new speciesof antbird from western Brazil. Wilson Bulletin 102:571- Grailariaridgelyi is oneof a numberof species 580. with very restrictedranges along the middle LOWERY,G. H., ANDJ.P. O'NEILL. 1969. A new spe- Rio Marafi6n,and as suchit is potentiallyvul- ciesof antpittafrom Peruand a revisionof the nerable to habitat alteration and other unnatu- subfamilyGrallariinae. Auk 86:1-12. ral disturbances. Hence, 700 ha of forest sur- LOWERY,G. H., ANDD. A. TALLMAN.1976. A new ge- rounding the type locality of G. ridgelyiwas nus and speciesof nine-primariedoscine of un- purchasedin October1998 and turned into a certain affinities from Peru. Auk 93:415-428. reservelarge enough to hold a viablepopula- MAYR,E., ANDE VUILLEUMIER.1983. New speciesof tion of the antpitta.The reserveis ownedand birds described from 1966 to 1975. Journal ftir managedby "Fundaci6nJocotoco," a nonprofit Ornithologie124:217-232. organizationfounded with thepurpose of pre- O'NEILL,J. E, ANDG. R. GRAVES.1977. A new genus servingand promotingthe study of globally and speciesof owl (Aves:$trigidae) from Peru. Auk 94:409-416. endangeredbird speciesthrough purchase and O'NEILL, J.P., C. A. MUNN, AND I. FRANKE. 1991. managementof land in Ecuador.Grailaria ridge- Nannopsittacadachilleae, a new speciesof parrot- lyi also occurson neighboringland in the let from eastern Peru. Auk 108:225-229. southernend of PodocarpusNational Park, un- PACHECO,J. E, B. M. WHITNEY, AND L. P. GONZAGA. derscoringthe importanceof the park,which 1996. A new genus and speciesof furnariid already holds severalknown threatenedand (Aves:Furnariidae) from the cocoa-growingre- near-threatenedspecies (Rasmussen and Rah- gionof southeasternBahia, Brazil. Wilson Bul- bek 1994), in preservingthe diversityof An- letin 108:397-433. dean avifauna. PARKER,t. A. III, T. S. SCHULENBERG,G. R. GRAVES, AND M. J. BRAUN.1985. The avifauna of the

ACKNOWLEDGMENTS Huancabambaregion, northern Peru. Pages 169- 197in Neotropicalornithology (P. A. Buckley,M. We thank M. MorenoEspinosa (Museo Ecuatori- S. Foster,E. S. Morton, R. S. Ridgely, and E G. ano de CienciasNaturales) and S. Lasso (Ministerio Buckley,Eds.). Ornithological Monographs No. de Agriculturay Ganaderla)for help in obtaining 36. collectingand export permits; A.D. Brewer,J. and R. RASMUSSEN,J. E, AND C. RAHBEK.1994. Aves del Moore, X. Mufioz, R. S. Ridgely,M. Rivadeneira,N. ParqueNacional Podocarpus. Una listaanotada. Simpson,and D. L. Wechslerfor companionshipin C.E.C.I.A., Quito, Ecuador the field; M. Rivadeneira and X. Mufioz of Neblina RIDGELY,R. S., ANDJ. A. GWYNNE,JR. 1976. A guide Forestfor ablyhandling the complexlogistics of our to the birds of Panama.Princeton University field expeditions;Montgomery Sisam Associates Inc. Press,Princeton, New Jersey. Architectsfor useof theircomputer; and M. B. Rob- RIDGELY,R. S., AND G. TUDOR.1989. The birds of bins, L. Joseph,A.D. Navarro-Sigtienza,and D. E. SouthAmerica, vol. I. Universityof TexasPress, Willard for valuableguidance and usefulcomments Austin. on early draftsof the manuscript.Financial support RIDeELY,R. S., AND G. TUDOR.1994. The Birds of for our expeditionswas provided by J.and R. Moore SouthAmerica, vol. II. Universityof TexasPress, and N. Simpson.Curators at ANSP (R. S. Ridgely) Austin. and LSUMNS(E H. Sheldon)generously made fro- SCHULENBERG,t. S., AND M.D. WILLIAMS. 1982. A zen tissues available for research. S. Trammell of newspecies of antpitta(Grailaria) from northern ANSP drew the sketchfor Figure 1. Financialsup- Peru. Wilson Bulletin 94:105-113. 890 KRABBEET AL. [Auk, Vol. 116

SMITHE,E B. 1975.Naturalist's color guide. American VUILLEUMIER,E, ANDE. MAYR.1987. New speciesof Museumof Natural History,New York. birds described from 1976 to 1980. Journal ftir ULLOA,C., AND ?. M. JOERGENSEN.1993. Arboles y Ornithologie128:137-150. arbustos de los Andes del Ecuador Institute of BiologicalSciences, Aarhus University, Aarhus, Denmark. Associate Editor: R. M. Zink