COLOR PL^FE. ChapadaFlycatcher, new species tyrant-flycatcher a of (Tyrannidae: Suiriri) from the cerrado regionof centralSouthAmerica.

The male is pictured in profile on the left; the femaleis displaying with wingsraised,on the right. From a watercolor paintingby Daniel E Lane.

The Auk 118(1):56-78, 2001

A CRYPTIC NEW SPECIES OF FLYCATCHER (TYRANNIDAE: SUIRIRI)

FROM THE CERRADO REGION OF CENTRAL SOUTH AMERICA

KEVIN J. ZIMMER, ANDREWWHITTAKER, AND DAVID C. OREN 1'2'5 3 4

tLos Angeles County Museum NaturalHistory,900 Exposition of Boulevard, Angeles, Los California 90007, USA; 21665GarciaRoad,Atascadero, California93422, USA; 3Estrado Aleixo, do Conjunto Acariquara Ruada Samaumas Manaus, Sul, 214, Amazonas, Brazil69085;and 4Departamento Zoologia, de MuseuParaense Em'lio Goeldi,
Caixa Postal 399, 66.017-970 Beldm,Pard, Brazil

ABSTRACT.--A species tyrant flycatcher new of (Suiririislerorum) described is from the cerradoregionof Brazilandadjacent eastern Bolivia.Thespecies previously beenconfused had with Suiririsuiririaffinis, with which it is syntopic multiplesites.The new species at was firstidentified voice. by Although cryptically similarto S.s.affinis manyrespects, new in the species readilyidentified all vocalizations, size,colorpatternof thetail, andshape is by bill of the centralrectrices. Mostdistinctive the male-female are duets,whichare accompanied by dramaticwing-lifting displaysnot performedby any congeners. Reciprocal playbackexperiments tape-recorded of vocalizations demonstrated the new species S.s.affinis that and donot respond oneanother's to vocalizations. provideinformation thenaturalhistory We on of the new flycatcher, alongwith spectrograms its variousvocalizations. alsoprovide of We vocalanalysis all othernamedtaxa in Suiriri,and discuss variousintrageneric of the relationships. particular,S.s. affinis In and S.s. bahiae, althoughdistinctmorphologically, are vocallyand behaviorally similar,and respond oneanother's to vocalizations playback in experiments.Received December 27 1999,accepted September 5 2000.

bahiae a subspecies S. affinis, as of from which he foundit to differ from typicalformsby "uptionallywererecognized, short-billed, a white- pertail covertsdark hair brown like the tail; bellied nominate form Suiriri suiriri; and a rectrices withoutany yellowishat the baseand longer-billed, yellow-bellied form, S. affinis. without the pale brownishapicalband." The apparentintermediacy a large series of of Zimmer (1955) pointed out an additional Suiririfrom northeastern Paraguay Zimmer complication: AMNH specimens S.affinled five of (1955)to suggest that the two taxa were con- is from Mato Grosso and Goias, Brazil, all of specific. Meyer de Schauensee (1966)and Tray- which were anomalousin having a short, lor (1979)followedZimmer in recognizing only broad-based bill and with the "pale terminal onespecies Suiriri,whereas of Short(1975)did band on the rectrices unusuallywide and disnot agreethat Paraguayan birdsshowed signs tinct, exceeding any other specimen hand at of intergradation and continued to recognize whether from the same or other localities and twospecies. Traylor(1982) reexamined Par- whether suiriri, affinis, bahiae." the or Zimmer furaguayan materialin the American Museumof ther noted that "the significance this comof Natural History (AMNH), and found it as de- binationof characters completely is puzzling, scribedby Zimmer, concurringthat S. suiriri sincethere is no allied group toward which and S. affinisshouldbe treatedas conspecific. these features,singly or together,suggesta More recent authors have remained divided: trend." In his reexamination Suiriri, Traylor of Sibleyand Monroe (1990) recognizedtwo spe- (1982)commented the five aberrentAMNH on cies of Suiriri, whereas Ridgely and Tudor specimensand an identical specimen from (1994)recognizedonly one.As currentlytreat- Maranhao, Brazil (from the Field Museum of ed, the genusalsoincludes oneadditionaltax- Natural History,Chicago; hereafter FMNH). In on, S.s. bahiae, yellow-bellied taxon from additionto the shortbill and broad,pale tips to a northeasternBrazil. Hellmayr (1927) treated the rectrices, sixspecimens the furthercould be distinguished from typical affinis their disby E-mail: kjzsrz@tcsn.net tinctly broader central rectricesand a lack of
onomicdebatefor decades. Two species tradi57

THE GENUS SUIRIRI has been the focus of tax-

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ZIMMER, WHITTAKER, OREN AND

[Auk, Vol. 118

any contrasting pale edging to those feathers specimens and tissue sampleswere deposited (Traylor 1982). Traylor noted that "ordinarily with the MPEG. We also located additional the close correlation of two such discrete and sitesof sympatry betweenthe two vocaltypes.
unrelated characters as the short bill and the
METHODS

coloration and shapeof the rectrices would be strongevidence that we have two siblingspecies.I think that eventuallythis will proveto be true,but thepossibility thatthe short-billed birds may somehow relatedto the intergrabe dation betweenaffinisand suiriri cannotbe ignored at this time." He concluded stating by "most important,without field studiesof the
various taxa, it is uselessto speculate."
work at various sites in Brazil Zimmer and

We made observations

of Suiriri at various sites in

Mato Grosso,Brazil (August and October1991,and every Septemberfrom 1993 to 1999); Pernambuco, Brazil (each January from 1996to 1999,andFebruary 1996);Goias,Brazil (January 1996);andAmapa,Brazil (February1998and 1999).All measurements used in behavioraldata (e.g. distances, heights)are estimates. Sexual identification of birds in the field was

During the courseof severalyears of field basedon sex-specific, stereotyped displays vocalor
Whittaker foundthat therewere two markedly different vocal types of yellow-bellied Suiriri, oneof whichwasrelatively widespread, occurring from Amapa to Bahia to Goias;and the other which we knew only from the Chapada dosGuimaraes regionof Mato Grosso. first The type includedbirds that, by both distribution and morphological characters, clearlywereassignable S.s. bahiae, well asbirds that apto as peared to be typical of S.s. affinis. The second type was distinctivenot only for its vocalizations, but also for its dramaticwing-flapping displaysthat invariablyaccompanied terriall torial duetsby mated pairs (seebelow). Reciprocal playback experimentsconductedat multiple sitesrevealed thatthetwo vocaltypesdid not respondto one anothers'vocalizations. In September 1998,we foundthe two vocaltypes occurring syntopically near Cuiaba, Mato Grosso, Brazil.Closecomparison revealed that onevocaltype wasnoticeably short-billed comparedto the otherandhad a distinct buffy terminal fringe to the tail, and that the birdswere alwaysassortatively paired with their own vocal and morphological types. Suspecting the short-billed that birdsrepresentedthe samebirds as the anomalous specimensreportedby Zimmer and Traylor, we returned to the Cuiabaregionin September 1999
with Dionisio Pimentel of the Museu Paraense

izations,the sexualspecificityof which was confirmedby postmortem inspection specimens of that we collected. Mapped distributions theyappear (as in this paper) are basedentirelyon labeldatafrom specimens the seniorauthorexamined, more that on recentrecordsdocumented tape recordings, by and on our own collecting sites.Thoselocalities wereentered into a sector-based Geographic Information System(Isler 1997)and mappedby MortonIsler.
We assume that vocalizations of Suiriri, like those

of othersuboscines, mostlyor entirelyinherited are (Kroodsma1984,1989;Kroodsmaand Konishi1991), and as such,providepotentiallyinformative charactersfor systematicstudy as in other suboscines (Lanyon1978,Isler et al. 1997,Krabbeand Schulenberg 1997).To analyzevocalizations, assembled we recordings all namedtaxa of Suiririfor compariof sonwith our own recordings. Locations recordand istsfor all recordings examinedare listedbelow.For comparison, vocalizations were categorized loudas songs, duets,or calls."Loudsongs" wereconsistently patterned multinotevocalizations (Isleret al. 1997) givenby an individualbird, seemingly thecontext in of territorialadvertisement. Thosewere givenindividually by both sexes, mostfrequentlywere inbut corporatedinto duets. "Duets" involved simultaneoussingingby a pair of Suiriri, and consisted of typicalloudsongs both sexes, of combined with other calls.Vocalizations categorized callsmostoften as were structurally simple (typically involving only oneor a few notes; exceptions notedbelow)and are usually were given in the contextof contact vocalizationsbetweenmatesor family members.Our recordings were madewith SonyTCM-5000tape re-

Emilio Goeldi, Belem (hereafter MPEG) and corders and Sennheiser ME-80, ME-66, and MKH-70 microphones. Spectrograms were madeby collected specimens each 10 of vocaltype.Prior shotgun PhyllisIsler on a Macintosh computer G4 usingCato collecting, madevoucher we tape-recordings nary version1.2.1(Bioacoustics Research Program, of each bird (all recordings be archived the Cornell Laboratory of Ornithology, Ithaca, New to at Library of Natural Sounds,Cornell Laboratory York).Canaryparameters were standarddefaultexof Ornithology, Ithaca, New York; hereafter ceptthat frameoverlapusedwas>-96.88%. LNS). Tissuesamplesalso were preservedfor Playback experimentswere conductedto deterof molecular analysisof genetic differences.All minereactions Suiririof eachtype to vocalizations

January 2001]

NewFlycatcher Cerrado from Region

59

of the others.In eachcase,an individual or pair of flycatchers first was presentedwith a prerecorded tapeof anothertaxon.Eachprerecorded tapeincluded a combined10 to 15 songs and duets(interspersed with calls)from two pairs of birds of a given taxon and geographic locality.Forthe S.s. suiriritape,we usedonlysongs callsbecause had norecordand we ings of duets. For all trials, each taxon was representedby a singleplaybacktape consisting vocalof izationsof two pairs (or individuals in the caseof nominate Suiriri)of birds.Duringeach trial, thetape of 10 to 15 songs and duetswasplayedto conclusion. To avoid confounding effects, a buffer period of about2 min was usedto separate playbackof vocalizationsof differenttaxa.Flycatchers did not rethat spond playback othertaxathenwerepresented to of with a playbacktape of their own vocal type. That wasdoneto account potentialseasonal for influences by resolving question whetherbirdsthatwere the of unresponsive the first trials were discriminating in betweenthe differentvocalizations, were generor ally nonterritorial duringthe trial periodandthereforeunresponsive playback anykind. Failureto to of
account for seasonal differences in responsiveness

Museum of Natural History, YaleUniversity (YPM). A wing rule with a perpendicular stop at zero was used to measureflattenedwing chord (wing), tail length(tail), breadthof the pale terminaltip to the rectrices (pale tip), and width of the centralrectrix (centralrectrix), and dial caliperswere usedto measuretarsuslength(tarsus),culmenlengthfrom the anterior end of the nares to the tip (culmen),bill depth at the anterior end of the nares (bill depth), and bill width at the anterior end of the nares (bill width). All calipermeasurements weremadeto the nearest0.01mm, those taken with the wing rule
were made to the nearest 0.5 mm. General linear

models were used to investigategender-corrected differencesamong taxa for each of the characters measured.Residual diagnostics indicatedno violationsof the generallinearmodelassumptions erof ror normality and constantvariance.Plumage was describedfrom specimens and comparedto a standard color reference(Smithe 1975).
RESULTS

Examinationof our specimens and tape rehasbeencitedasa potentialdesignweakness othin of er playbackexperiments(Kroodsma1986).Respons- cordings,study of a large selection museum extensive field observations inthat es to playback were characterizedas "strong," specimens, "moderate, weak," or "none." A strongresponse cluded reciprocal playback experiments,and ....
involved immediate and repeated vocalizations (from previouslynonvocalizing birds) as well as ap-

vocalizing, or sustained vocalizing without approach. weak response A involvedsingleor unsustainedvocalizations (from previously nonvocalizing birds)withoutapproach. nonecategory The includes instances which in birds remainedsilent(in the case of previouslynonvocalizingbirds) and did not approach the sound source,as well as instances in whichalreadyvocalizing birds neitherchanged the delivery or rate of their vocalizations,nor approached soundsource. potentialfifth category the A wouldbe a "negativeresponse," whichthe subject in bird responded the soundstimulusby movingfarto ther away.No negativeresponses playback to were
observedfrom any taxa. Zimmer examined representative specimensof

discovery multiplesitesof sympatrywithout of evidence interbreeding of convinced thatthe us proach toward the sound source.A moderatere- two vocal types of yellow-belliedSuiriri repsponse involvedeithera cautious approach without resentdistinctbiologicalspecies. Furthermore,
it is clear that the short-billed form that we collected from Mato Grosso is the same unnamed

form that was first pointed out by Zimmer (1955)and later hypothesized be a cryptic to species Traylor (1982).We proposeto name by this flycatcher
Suiriri islerorumsp. nov.

ChapadaFlycatcher
Holotype.--MPEG No. 54867, adult female from kilometer 2 on the road to Agua Fria, 1525'S, 5546'W, westof the town of Chapada

Suiriri (n = 170) for comparison with our 20 speci- dos Guimaraes, approximately 720 m elevamens from Mato Grosso.Those specimensare tion, state of Mato Grosso, Brazil, collected 21
housed at the Museu Paraense Emilio Goeldi, Belem,

Brazil(MPEG);Academy NaturalSciences Philof of adelphia,Philadelphia(ANSP);CarnegieMuseum, Pittsburgh (CM); Field Museumof Natural History, Chicago(FMNH); Los AngelesCountyMuseumof Natural History,LosAngeles (LACM); theLouisiana
State University Museum of Natural Science, Baton

September 1999by DionisioPimentel,tape-recordedby Kevin J. Zimmer. Voicespecimento be archived at Library of Natural Sounds (LNS), CornellLaboratory Ornithology, of Ithaca, New York.

Diagnosis.--Similar plumage and size to in

Rouge (LSUMZ);theMuseum Vertebrate of Zoology, Suiriri s. affinis(Tables1 and 2), but differs as bill distinctlyshorter, relatively broadBerkeley(MVZ); the National Museum of Natural follows: History,Washington, D.C. (USNM); andthePeabody er, and more nearly uniform blackish (baseof

60

ZIMMER, WHITTAKER, OREN AND

[Auk, Vol. 118

TABLE Measurements(mm) of Suiriri islerorum 1. and congeners. Valuesare means _+standarddeviation followedby ranges,with samplesizein parentheses.

Characters
Culmen

S. islerorurn
8.0 _+ 0.7

S. affinis
9.6 -+ 0.6

S. bahiae
8.5 0.7

S. suiriri
7.7 _+ 0.4

Bill depth
Bill width

7.5-8.6 (23) 4.2 _+0.3 3.7-4.9 (23)

5.7 0.3

8.2-11.1 (61) 4.6 _+0.3 3.8-5.4 (61)

5.4 _+ 0.4

7.9-9.2 (3) 4.5 _+0.9 4.5-4.6 (2)

5.5 0.1

6.7-8.7 (108) 4.2 _+0.3 3.7-4.9 (106)

5.0 0.3

Wing
Tail

5.1-6.3 (23) 82.6 _+3.3 72-87 (21)

71.5 2.7

4.7-6.1 (61) 82.0 _+4.4 70-88.5 (38)

69.4 + 3.8

5.4-5.6 (2) 77.2 4.0 73.5-81.5 (3)

66.7 4.2

4.4-5.6 (107) 73.0 3.3 65.5-81 (108)

69.0 3.6

66-78 (21)
Tarsus 20.3 + 0.7

59-76 (38)
20.5 + 1.0

62-70 (3)
20.3 0.7

58-78.5 (108)
19.8 _+ 0.7

Pale tip
Central rectrix

19.1-21.7 (20) 6.4 + 1.5 3-11 (23)

10.5 _+ 1.1

17.2-22.5 (37) 2.1 1.5 0.0-4.8 (61)

8.6 1.3

19.8-21.1 (3) 1.2 2.0 0.0-3.5 (3)

7.0 + 1.0

17.9-22.1 (106) 0.0 - 0.0 0.0 (108)

--

8-12 (21)

6-11 (38)

6-8 (3)

--

mandiblefleshy-pink manyspecimens afin of finis, particularly in females);dorsalsurfaceof

S. islerorurn; to the bottomof thebill in all only S.s. affinisexamined).Suiriri islerorurn differs most of tail darker, more blackish-brown further from specimens S.s. bahiae having of in (browner, between Fuscous,color#21, and Van- the grayish crown and nape less sharply dedyke Brown, color#121, in affinis); pale termi- marcatedfrom the greenerback;back darker nal fringeto tail broaderand morehighlycon- and less distinctly green;breastlesswhitish; trasting (either absent,or, if present,narrow rump and uppertail covertsmorecontrastingly and weaklycontrasting affinis); in both websof pale; and central rectricesbroader and not differs from S.s. outerrectrices more extensively pale, and dark pale-edged. Suiriri islerorurn subterminal patchon innerweb of outerrectri- suiriri as follows:distinctlylarger overallwith cesdistinctlysmaller;centralrectrices broader proportionately shorter, broader tail; belly, and without paler edges;white of throat ex- flanks,thighs,and crissum yellow ratherthan tendsfartherup on sides face(to the gapein white; undertailwith a broad,contrasting of pale
TABLE Means SD (samplesizesin parentheses) measurements 2. of (mm) of Suiriri islerorum S.s. and affinis sex,followed a statistical by by comparison themeans thepooled of of samples (bothsexes) each for species. Valuesof P (= probabilityof a greatervalueof F) derived from Analysisof Variance(ANOVA).
islerorum
Characters
Culmen

affinis
Males
9.8 -+ 0.5

Males
8.1 0.4

Females
8.0 0.3

Females
9.3 0.5

P
<0.0001

(6)

(16)

(31)

(26)

Bill depth
Bill width

4.2 0.4 (6)

5.8 0.4

4.2 0.3 (15)

5.6 0.2

4.7 0.3 (31)

5.5 0.3

4.6 0.4 (26)

5.3 + 0.4

<0.01
<0.01

(6)

(15)

(31)

(27)

Wing chord
Tail

85.1 0.7
(5)
71.2 1.6

81.6 3.6
(14)
71.0 _+ 2.7

85.0 _+2.3
(lS)
70.9 2.9

78.6 3.8
(17)
67.8 _+ 4.1

<0.05
<0.01

(5)
Tarsus 21.2 0.6

(14)
20.0 ___ 0.06

(lS)
21.2 0.6

(17)
19.7 + 0.8 NS

Pale tip
Central rectrix

(5) 6.9 0.8 (6)

9.6 1.1

(14) 6.2 1.7 (16)

10.9 0.9

(17) 2.2 1.5 (31)

8.8 _+ 1.2

(17) 2.2 + 1.6 (27)

8.3 + 1.3

<0.0001
<0.0001

(5)

(16)

(lS)

(17)

January 2001]

NewFlycatcher Cerrado from Region

61

fusinginto a grayer hindcrown nape,closand est to Glaucous (color#79).The grayishnape

gradesinto an Olive (color#30) backand scapulars. Uppertail coverts and rump contrastingly paler than back,betweenLight Drab (color #119C) and SmokeGray (color#45). Auriculars

concolorwith hindcrown,perhaps slightly duskier Malar region between whitish and

Pale Neutral Gray (color #86). Lores slaty. A narrow,pale, supraorbital brow extendsto the forehead. Fourlongrictalbristles bothsides on of gape,the longest8.0 mm. Wingsgenerally dusky,closestto VandykeBrown (color#221), but secondaries narrowly pale-edged(closest to Light Drab, color#119C)on outerweb.Basal portion of inner web of all remiges broadly pale-edged,betweenPaleHorn (color#92) and FIc. 1. Displaying pair of Suiriri islerorum: female in foreground with wingsraised. Keymorphological Drab-Gray (color #119D). Greater and median covertsbroadly tipped Light Drab features distinguish species that this from S.s.affinis secondary can be seenin this photo:the short, thick bill, and (#119C),forming two distinctwingbars.Axilthe broad,pale, terminal fringe to the tail. Photo- lariesand underwingliningspale SulphurYelgraphed nearCuiab,MatoGrosso, Brazilon22Sep- low (color #57).
tember 1999by Kevin J. Zimmer.

Chin and throatwhitish,gradingposteriorally into Pale Neutral Gray (color#86). Center and sidesof breastGlaucous(color#80), separatingwhiter throatfrom SulphurYellow(color #57)belly,flanks,thighs,andundertailcoverts. Most of exposeddorsal surfaceof tail dark, between Dusky Brown (color #19) and Dark

base (undersideof tail in S.s. suiriri entirely dark except contrastingly for palenarrow outer web of outerrectrix on eachsideand pale shaft of outer rectrices);central rectricesdistinctly broaderandnot narrowlypale-edged either on web; inner web of outer rectrix pale-based and pale-tipped, entirelyblackish; not broader gray band acrossthe breast;less extensivelydark auriculars; lessdistinctwhite supraorbital and
brow. Suiriri islerorum differs from both Suble-

GrayishBrown (color#20). Contrasting paler

band at tips of rectricesbetween Drab (color #27) and Dark Drab (color#119B); that palertip

7 mm in lengthon central rectrix.Base both of

webs of outer rectrices, and base of inner web

gatusmodestus and Sublegatus arenarumas follows: markedly larger; greener upperparts showingmore contrast betweennapeand back (brownerwith little or no contrastin Sublegatus);rump and uppertail covertscontrastingly paler than back (no contrast Sublegatus); in uppersideof tail blacker(lessbrown) with a pale

of all otherrectrices contrastingly pale,closest to Cream color (#54). That paler baseto tail
mostly concealed from aboveby uppertail coverts, but visible from below as a wide (35 mm) pale base that contrastswith a dark central panel (closestto Vandyke Brown, color #221), whichin turn, contrasts with the paleterminal

base therectrices; to throatcontrastingly white. The new species differsfrom mostmembers of Elaenia lacking elongatecrown feathersand in a contrastingly colored, concealed coronal patch, well asin havingan entirely as blackish bill. The contrastingly pale rump and baseof
the tail are characters not found in Elaenia.

fringe.Basalportionof the shaftsof mostrectrices are closestto Pale Horn (color #92); those
of the central rectrices are closer to Salmon

(color #6). Outermost rectrix on each side has the outerweb almostentirelypale, with a dark

subterminal patch(26mm in length)on theinner web.Thepaleouterwebof theouterrectrix, with the palebaseto the rectriDescription ofholotype.--See Figure 1 and col- in combination or plate. Capitalized colornamesand numbers ces and the pale terminal fringe, isolateand are from Smithe (1975). Forecrownbrownish outlinethe dark central panelon theunderside gray,closest Hair Brown (color#119A), suf- of the tail. to

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ZIMMER, WHITTAKER, OREN AND

[Auk, Vol. 118

Tocantins, and Maranhao. The westernmost

records from Campodo Estanho, are Amazonas, Brazil (M. Cohn-Hafttape recording),and ParqueNacionalNoel KempffMercado,depto.
Santa Cruz, Bolivia (LSUMZ 150871). Un-

doubtedlyoccursmore extensively within the area boundedby thoserecords,but probably overlooked to morphological due similarityto
S.s. affinis.

Elevational distribution.The only specimens

of Suiriri islerorum with elevational data on the labels are the ones that we collected in Mato

FIG. 2.

Distribution

of Suiriri

islerorum

and its

Grosso, Brazil. Thoserangefrom 250 m on the Coxipodo Ouro RoadnearCuiaba,to 720m on the Agua Fria Road near Chapada dos Guimaraes. That elevationalrange also encompasses of our records all that havebeendocumented by tape recordings. The Bolivian specimen(LSUMZ) was from 450 m (J. Bates pers. comm.).Paynterand Traylor (1991)provide elevations for several of the sites from

congeners confirmed examination specimens as by of or tape recordings: blackcircles= S. islerorum; diamonds= S.s. affinis; circlestouchingdiamonds= sites where islerorum and affinis are sympatric; squares S.s.suiriri;plussigns= sites = whereaffinis and suiriri are sympatric; and triangles= S.s.bahiae. "T" next to a symbolindicates type localityfor the
that taxon. "T" next to a circle-diamond locates Cha-

which S. islerorum were collectedby other workers. Thoseshouldbe viewedaspertaining to generalcollecting localities, and may not apply specifically siteswhereSuiririwere colto lected.Includingthosedatawould extendthe probableelevationalrange of S. islerorum from
150 m at Tocantinia, Tocantins, Brazil (LACM 46004) to 1200 m at SaoJoaoda Allanna, Goias, Brazil (LACM 40087). Specimens examined: Skins.nly specimens

pada dos Guimarfies, Mato Grosso, Brazil,the type localityfor S. islerorum. Note that no type localityis
located for S.s. bahiae,which was described from a

trade specimen whose only locality was "Bahia." Map by Morton Isler.

Soft parts in life: iris, medium brown; maxilia, black; mandible, basal one-quarter slaty

measuredare listed.Plumages all sympatric of speciesof Elaeniaand Sublegatus were compared superficially. Suiriri islerorum: Brazil,
Mato Grosso, six males and nine females

gray,remainder black; lining of themouth, yellow-orange; tarsusand toes,black;solesof feet and toes,paler gray;nails,black. Measurements holotype.--Culmen of from anterior edge of nares,7.8 mm; bill depth at anterior edgeof nares,4.4 mm; bill width at an-

(MPEG 54865 to 54874, and 19299; AMNH 127882,127883,33101, and 33135);Gois, three females (LACM 460087 and 40089, MPEG

19299);Maranhao,one female (FMNH 63435); Para, one female (MPEG 49036);Tocantins,one female (LACM 46004); Mato Grossodo Sul, one (ANS 159808). Bolivia,depterior edge of nares, 5.8 mm; wing chord, unsexedspecimen to. Santa Cruz, one female (LSUMZ 150871).Of 85 mm; tail, 73 mm; tarsus, 20.1 mm; body mass22.8 g. Ovary, 6 x 3 mm; oviduct,3 mm those,four malesand six females(MPEG 54865 to 54874)were collectedby us in Mato Grosso, wide; largestovum, 4 mm. Geographic distribution.--Locally distributed Brazil. Suiriri suiriri affinis: Brazil, Amap, 10 males in cerrado and camposhabitats throughout and 9 females (MPEG 46607 to 46619, AMNH much of central Brazil, south of the Amazon and eastof the Rio Madeira,and at leastspar- 33100, USNM 515199);Goias, five males, six fe(LSUMZ ingly to depto.SantaCruz, easternBolivia (Fig. males and two unsexedspecimens 2). Most confirmed records are from Mato 31774, 67522, 67523; ANS 186661; USNM

Grosso, Brazil, with scattered specimens east

and north to the Brazilian states of Par, Gois,

516085; MPEG 22013; FMNH

344593 to 344595

and 75099; LACM 46003, 40090, and 40088);

January 2001]
Mato

NewFlycatcher Cerrado from Region

63

Grosso, seven males and six females

providedin the corresponding figurelegends.

All recordings Kevin J. Zimmer unlessothby
erwise indicated. Suiriri islerorum: Brazil, Mato

(MPEG 54875 to 54884, AMNH

33100, 33131,

and 33133); Maranho, one male and five females (FMNH 63431, 63432, 63434; MPEG
43526 to 43528); Minas Gerais, one male and two females (MPEG 36210, FMNH 56657,

Grosso, westof Chapada Guimaresalong dos the roadto AguaFria, 20 specimens, Coxand

ipo do Ouro road near Cuiab, 8 specimens; Amazonas,160 km E-SE of Humaita, 1 specimen (M. Cohn-Haft).Suiririsuiririaffinis: Brazil, Amap, 12 specimens; Gois, 2 specimens (A. Whittaker); Mato Grosso, westof Chapada dosGuimaraes alongthe road to Agua Fria, 4 specimens, Coxipodo Ouro road near Cuand iab, 24 specimens. Bolivia, depto.SantaCruz, Suiriri suiriri bahiae:Brazil Bahia, one male 1 specimen(T. A. Parker,LNS 52044). Suririri Brazil, Pernambuco, near Lagoa and two females (LACM 37067 and 37068, suiriri bahiae: Grande,16 specimens. Suiririsuiriri suiriri:ArFMNH 64119). (S. Suiriri suiriri suiriri: Argentina, Entre Rios, gentina,Salta,three specimens Hilty). Boseven males, five females, and one unsexed livia, depto.SantaCruz, one specimen A. (T. specimen(YPM 66068 to 66073 and 83326, CM Parker,LNS 33629).Brazil, Rio Grande do Sul,
LACM 40086); Tocantins, one male (LACM 46005);Cear, one male (USNM 264644);PiauL one unsexedspecimen(MPEG 50917).Bolivia, depto. Santa Cruz, three males (LSUMZ 124634, 124642, 150872). Of these, six males and four females(MPEG 54875 to 54884) were collected us in Mato Grosso,Brazil. by
140482, 140489, 140490, 140670, 140806, and

one specimen (W. Belton, LNS 19531). Para-

140867);Misiones, five males and five females (LSUMZ 56714 to 56723); Tucuman, three

guay,Chaco Province, specimen Finch, one (D.

LNS 57891).

males,four females,and two unsexed specimens (FMNH 58012, 58019, 58369, and 58377; USNM 284898, 285001, 285002, and 285109;

Etymology.--We take great pleasurein nam-

ing thisspecies afterour goodfriendsandcolleagues, Morton and PhyllisIsler, in recogni-

YPM 24195);La Pampa,one male and two fe- tion of their numerous contributions to Mort and Phyllisaumales (USNM 284329, 284332, and 284335); Neotropicalornithology. on Chaco, one male and one female (USNM thored a landmark monograph tanagers 284330 and 284334); Buenos Aires, two males (Isler and Isler 1987),and they havemadepicontributions developing in method(USNM 55681and 55683);Salta,onemale (CM oneering for of to 45839); and Santiago del Estero, one male ologies the analysis vocal characters species limits in antbirdsand othersub(MVZ 108323). Bolivia, Santa Cruz, 18 males, assess 14 females, and 1 unsexed specimen(CM oscines(Isler et al. 1997, 1998, 1999). Their 119820, 120212, 120213, 32786, 32787, 32831, work on antbirds served heighten has to aware32889, 32891, 32952, 51402, 78970, 79244, nessof the importanceof vocalizations taxas 80363, and 80364; FMNH 181493, 181502, onomic characters all suboscine in groups, and
181506, 294367, 295413, 296295, 296298, 296301, 335151, and 335152; LSUMZ 124633 to 124637, 153757, and 153758; YPM 31816 and

as such,has far-reaching implications the for

accurateassessment regional endemismand of biodiversity,and, ultimately,for conservation.

greatlyto our own 31817); Tarija, five males and three females The Islershavecontributed birds, and are constant (ANS 138951, 138952, 138954, 138955, 138958, studiesof Neotropical and help. and 138959; FMNH 294369 and 294370); Co- sourcesof advice, encouragement, namecalls attention thetypeloto chabamba, one male and two females(FMNH TheEnglish 181498 and 181500;ANS 136168). Paraguay, cality,theChapadadosGuimares,Mato Grosof depto. Presidente Hayes, two males (MVZ so, Brazil, whichis an importantreservoir 168120and 168121);depto.Chaco,four males threatened cerrado flora and fauna.
(FMNH 153007,153010to 153012).Brazil Mato
Grosso do Sul, one male and two females
REMARKS

(FMNH 64121 to 64123); Parana, one male and

onefemale(USNM 16367and 16368). Uruguay, Sexual dimorphism variation thetypeseand in RepresaPalmar,one male (ANS 187738). ries.--The type seriesof Suiriri islerorum conSpecimens examined: Tape recordings.--Data sists 23 adult specimens males,16females, of (6 for recordings reproduced as sonogramsare and 1 unsexedspecimen). Males resemble fe-

64

ZIMMER, WHITTAKER, OREN AND

[Auk, Vol. 118

males, and plumage variation among paratypesappearsto be neithersexuallynor geographically influenced. There is some individual variationin colorof the uppertail covertsand in the extentof the contrastingly palebaseof the rectrices. Compared thehoto lotype, someindividuals (MPEG 54872,LACM
46004, LSUMZ 150871, AMNH 127882) are

wear. We believe that all of those birds were in

breedingcondition.We neverhave observed

family groupsof S.islerorum during ourAugust

and September visits to Mato Grosso.That, combinedwith the apparentbreedingreadiness and territorial nature of the birds we col-

more greenish-yellow (closest Citrine,color to #51) on the uppertailcoverts, another(MPEG

54870) was buffier, and two (MPEG 54871 and

FMNH 63435) had theuppertailcoverts nearly the same color as the back. Two specimens
(MPEG 54865 and LACM 40087) are more ex-

lectedin lateSeptember, suggests themain that hatching fledging and periodis probably Ocin tober.That would fit a common patternfor insectivorous birds in regionswith pronounced dry seasons, whichhatchingcoincides in with onsetof the rainy season and the concomitant flush of insect populations(Walsberg1977,
Marr 1981, Zimmer 1993).

The four AMNH specimens from Mato Grostensivelygray dorsally,with the mantle and lowerbackmorenearlyconcolor with thenape. so were collectedbetween 7 February and 26 Similar variation in rump-lower-backcontrast May. All had remigesand rectrices that were was apparentamongthe 61 specimens S.s. less worn and more broadly pale-eged or of fringed than in our specimens. Similarly,the affinisexamined. from Gois and TocanMost observedvariation among paratypes threeLACM specimens
was related to feather wear and molt condition.

tins were collected between14April and3 May,

plumage thanourseries Most specimens browneron the forecrown andall werein fresher are The (ANSP and grayer on the hindcrown,but a few are from Mato Grosso. ANSP specimen moreuniformlygraythroughout crown.In- 159808)was collected30 Juneand was in fresh the dividual feathers of the forecrown have a condition. The LSUMZ specimen (LSUMZ brownishlongitudinalstreakalongthe shaft, 150871) was collected29 August, and was in and are gray aroundthe fringe. When those fresh condition with a trace of molt on the head 6 feathers worn,theextentof thegrayrelative and with an ovary that measured x 7 mm. are to the brownish central streak is reduced, and Thecondition those of specimens suggests that the forecrown assumes morebrownishtinge. S. islerorurn a undergoes one molt sometime after Freshly molted birds had grayer forecrowns. breedingin September October,and then to Similarly,variationin the breadth of the pale probably undergoes at least a partial molt terminal fringe to the rectrices, buffy tips again in spring or summerprior to the next the of the secondary coverts, thepaleedges and of breedingseason. Vocalizations.--Male and female Suiriri islethe remigesall wasinfluenced wear. by One male specimen(MPEG 54871) had an rorum havedifferentloudsongs, whicharetypunusuallylong tip (hook)to the maxilla.The ically delivered as sex-specific elements siof culmen measurement for that bird was 8.52 multaneous duets, but are occasionally mm, but without the tip the culmen would deliveredseparately. The male loudsong (Fig.
measure 7.64 mm. In general, males were 3A) is a loud seriesof paired couplets, notes all

slightlylargerthanfemales, although mea- having a distinctlytwangy quality.The song bill surements were nearly identical(Table2). hastwo discernible phrases arerepeated that in Breeding molt.--Tenof the 23 specimens predictable and patterns(where where, whooz whooz it
of Suiriri islerorum (MPEG 54865 to 54874) were
collected from Mato Grosso, Brazil between it, where where whooz it, whooz it, whoozit, whooz

it). Sometimes males will sing a series of

"whooz it" notes without the "where where" notes. The number of consecutive "whooz it"

20-22 September 1999. All had enlargedgonads, and two females (MPEG 54866 and

54872)had well-developed broodpatches. All

were vocal and territorial. Most birds showed

couplets may vary from two to six (rarely more), but no more than one"where where" cou-

signsof moderate heavywearof the secondto ary coverts rectrices. and Similarly, FMNH the specimen from Maranhfio wascollected Sep2 tember 1925, and showedmoderateto heavy

plet seems be givenwithoutinterjecting to the "whooz it" notes. In responseto playback, malesfrequentlysinglongersongs. The most commonlyheard male calls are isolatednotes

January 2001]

NewFlycatcher Cerrado from Region

Suiriri islerorum
B calls

65

MALE

A loudsong
4

3
2

'
o

1
i i i i i i i i i

0.5
FEMALE

I .0

I .5

2.0

2.5

3.0

3.5

4.0

4.5

5.0

"'

C loudsong

D "wheer-deer"

E "zhuwheep" "zhuwheep-oo"

..

0.5

I .0

1.5

2.0

2,5

3.0

3.5

4.0

4.5

5.0

Time

(seconds)

FIG. 3. Vocalizationsof Suiriri islerorum recordedfrom Mato Grosso(MT), Brazil. MPEG numbersin pa-

rentheses matchtape recordings with specimens the MuseuParaense in EmilioGoeldi,Be16m, J, Brazil. Par (A) Male loudsong (MT, near Cuiab& 22 September 1999).(B) Male (MPEGno. 54874)calls(MT, Chapada
dos GuimarSes,21 September1999). (C) Femaleloudsong(MT, near CuiabJ, 22 September1999).(D) Female

(MPEG no. 54872)"wheer-deer" (MT, Chapadados GuimarSes, September call 21 1999).(E) Female"zhuwheep" and "zhuwheep-oo" (MT, near CuiabJ,22 September calls 1999).All recordings Kevin J. Zimmer. by

or couplets similarto thosegivenin loudsongs ble of notes, with the female rattle and the (Fig. 3B). The femaleloudsong (Fig. 3C) is a twangy male notesequallydistinct. As with S. islerorum, the male and female loud,bubblyrattleof variable lengththatis typically precededby one or two well-differenti- loudsongs S.s. affinisare most commonly of ated "whur" or "wheer"notes.In response to heardas sex-specific elements simultaneous of playback, femalestypically lengthenthe rattle duets,but are sometimes deliveredseparately. of is element their songs. of Females commonly give The male loudsong S.s. affinis a seriesof "pi-chew" notes, variable length,that in two types of calls. The first is a seriesof sneezy "wheerDEER" (Fig.3D) thataresomewhat typically begins with severalsqueakyintrocalls similarto the male couplets, thatare more ductory notes and ends with severalone-sylbut emphatic (particularlyon the second syllable), labled"chew" notes(Fig.5A). The femaleloudwith a lesstwangy,lessstridentquality, which song(Fig. 5B) is a series loud, squeaky of notes arefrequentlyinterjected into duets.Outsideof thattypicallydecelerates noticeably towardthe duets,the mostcommonly heard femalecall is end. Thosenotesare similar in quality to the a querulous sounding,two-or-three-syllabledintroductory notes malesongs, arehighin but "zhuwheep" "zhuwheep-oo," or in whichthelong er frequency,higher amplitude, and more The terminal notesin a female first note is diphthongaland distinctlyup- widely spaced. ward-inflected at the end, and the second, loudsong often drop in frequency. Females shorter note,whenpresent, dropsin frequency have a variety of single-notecalls, most of (Fig.3E).That call is typicallygivenin thecon- which are nasal and have a distinctlywhiny text of a contact call when a femaleis visually quality(Fig.5C,D). Those routinely are uttered calls, whena female but gives several separated from her mate.In duets(Fig.4), the ascontact maleand femalesingtheir loudsongs simulta- differentiated whiny callsin rapid succession swiroo, chew")it is usually the neously, creatinga somewhat discordant jum- ("suiree,swee,

66

ZIMMER, WHITTAKER, OREIN AND

Suiriri
lO

[Auk, Vol. 118

islerorum

8 6
4 2 o 0

male/femaleduet

male/female duet

10.0

8 C male/female duet
!

.. ;,
i i

'...',. :...,
i i i

.... ....
i

..............
i i i

1.0

2.0

3.0

4.0

5.0

6.0

7.0

8.0

9.0

10.0

Time

(seconds)

FIG. 4.

Male-female duets of Suiriri islerorum recorded from Mato Grosso (MT), Brazil. MPEG numbers

in parentheses matchtape recordings with specimens the Museu Paraense in Emilio Goeldi, Be16m, Parfi, Brazil. (A) male-female (MPEG nos. 54865and 54866)duet (MT, near Cuiabfi,20 September1999).(B) Malefemale (MPEG nos. 54874 and 54872) duet (MT, Chapada dos Guimarfies,21 September1999). (C) Malefemale(MPEG nos.54869and 54867)duet (MT, Chapadados Guimarfies, September 21 1999).The female from this pair (MPEG no. 54867)is the holotypefor the species. recordings KevinJ.Zimmer. All by

synchronizing thatbegins duet.A female, cue a whosemate we had collected, repeatedthose single-note nasal calls for more than an hour afterwards. The most frequently heard male call outsideof a duet contextis a single-note, somewhat guttural "werk" (Fig. 5E). Lesscommonly heardis a low-frequency rattle that may functionas an aggression call (Fig. 5E G). We have heard that call given severaltimes from birds of unknown sex within foragingfamily groups,as well as from a female that was attempting to aggressively displace a Lesser Elaenia (Elaeniachiriquensis) from a fruiting tree.In duets(Fig. 6), the higheramplitudefemale loudsongs tend to mask the male loud-

The male loudsong S.s. bahiae of beginswith a rapid series squeaky, of somewhat run-together notesthat drop in frequency beforeyieldingto a distinctive series of "pitty-chew" elements (Fig. 7A). The introductoryseriesrecallsthe sounds that would be madeby squeezing toy a
rubber duck severaltimes in rapid succession.

The female loudsong(Fig. 7B) is a seriesof squeaky "choop" notes (peak frequency of
about 8 kHz) delivered at about 8 to 10 notes

songs. similarityof thefemalenotesto the The

introductory male notes makes it even more difficultto distinguish full male song. the All vocalizationsof S.s. bahiae appear to be

homologous the vocalizations S.s. affinis. to of

per second. Thosenotesare similar in tonality to thosethatprecede "pitty-chew" the elements in the male song,but are higher in frequency, more widely separated, and do not drop in frequency through series. the Bothsexes givetheir respective songs during simultaneousduets (Fig. 6D). In duets,the femaleloudsong typis ically louder than that of the male. A single duet bout might include two or more songs from eachbird, with the final female songin

January 2001]

New Flycatcher Cerrado from Region

67

perchin the crownof a tree the bout often decelerating noticeably toward sumeda prominent the endof the song.Theterminalnotesin such or on a baresnag,usuallywithin 0.5 m of one by a decelerated series lowerin frequency are than another Duets were always accompanied the introductory notes.Males frequentlycon- dramatic, sex-specific, stereotypedmotions. cludeduetswith a few sputtering "pitty-chew" The male flickedboth wings high abovehis calls. Outside of duets, the most frequently back (the wings nearly touching)at a rate of heard male call is a guttural, singlenote (Fig. about one per secondfor the duration of the 7C). Thiscall apparently functions a contact duet (6-10 s). With each downstroke of his as call.The series "pitty-chew" of callsalsois fre- wings,he flippedhis tail upwards,at abouta quently givenindependent the squeaky of pre- 45 angle.When the wings were once again lude (Fig. 7D), particularlyin duets.The most flickedupwards,the tail was flipped down 30 During thedisplay, commonly heard femalecall is a nasal"nyew" to 45belowthehorizontal. with a distinctly whinyquality.Those callsare the maleleanedforward,sothat his rump was typicallydelivered singlyin thecontext con- held at aboutthe sameplane as his head. The of tactcalls, theyalsoareoftenstrungtogeth- tail wasfannedthroughout but much thedisplay, of er in rapid succession (Fig. 7E), particularly by emphasizing contrast the betweenthe yellowbirds agitated by tape playback. Femalesre- ish-buff base and the blackish distal half. Some spondingto playback typically begin giving males rotated at approximatelya quarter turn thosewhiny notesin flight as they approach with each up-and-down cycle thewingsand of thesound source. in S.s.affinis, rapidlyde- tail, suchthat they wereconstantly As a turningin livered seriesof thosefemalenotesfollowing circles whiledisplaying. Simultaneous the with display, femalealsoraisedherwings the playback appears be the synchronizing to cue male's thatprecipitates responsorial fromthe high above back,but heldthemthere,fullythe duet her pair The only vocalization S.s. affinisfor fanned, for about 2-3 s at a time before lowerof whichwe couldfind no homolog S.s. bahiae ing themgradually. in Females repeated that mowasthe rattlecall (Fig.5EG).Theapparent ab- tiontwo or threetimesduringeachduet.At the senceof sucha vocalizationin S.s. bahiae may sametime that a femalewasraisingher wings, be an artifact of sampling,because rattle calls shewould alsofan and dip her tail downward were only occasionallyrecorded from S.s. at a 20 to 45angle,flippingthe tail backup to the horizontalpositionasthe wingswerelowaffinis. Our inventory of tape recordingsof nomi- ered.The fanningof the wingsby females acnate S.s. suiriri is small.What appearsto be a centuated the largely pale inner webs of the loudsong Argentina (Fig. 8A) is a long (3 to remiges such that backlit birds appeared to in wings.Females frequentalso 5 s), frenetic,sputteringseriesof nasal notes havetranslucent that riseand fall in frequency, with occasional ly rotatedwhile displaying, wereless but likely differentiated querulous notes scattered to do so than the frenetic males. Females also throughthe middle. We do not know if this maintained moreuprightposture a throughout Thosedisplays S.islerorum of acloudsongis specificto one sex, and, if so, to their displays. which one. The most common vocalization is a companiedeach of the more than 200 duet and are remarkably singlenasal"row?'or "jyow"note that is re- boutsthat we witnessed, Tyrant peatedincessantly prolongedperiods.Mi- similar to thoseof the Streamer-tailed for nor variations of this call lave been recorded (Gubernetes yetapa) southern of SouthAmerica, Flyfrom Argentina,Brazil, Bolivia, and Paraguay and, to a lesserextent,of White-bearded (Phelpsia inornata) Venezuela Zimof (K. (Fig. 8B-E). Anothercall recorded from Para- catcher guayis a 2-3 s rattle at about5-6 kHz (Fig. 8F). mer pers. obs.). We never have witnessedsimilar displays Duetdisplay behavior.--The moststrikingbehaviors of S. islerorum were those associated from eitherS.s. af, or S.s. bahiae, spiteof finis in more than 200 duetsof the with territorialduets matedpairs(Fig.1 and having witnessed of colorplate).Pairswere tightly synchronized in former and more than 100 duets of the latter. their duets. The first few notes of either a male Sick(1993)described songof S.s.affinis the as or female loudsongwere enoughto initiate "repeatedwhile lifting bothwingsvertically." mostduets,with the second bird joiningin im- That description almostcertainly based is upon of Suiriri s. af, finisand mediately.Prior to singing, pairs typically as- observations S. islerorum.

68

ZIMMER, WHITTAKER, OREIN AND

[Auk, Vol 118

Suiriri suiriri affinis

54-

A male loudsong
...

. intro

"pl-chew" elements

"chew" notes

321

0 0

I .0

I ,5

2.0

2.5

3.0

3.5

4.0

4.5

5.0

10
987-

B female loudsong

6 5
4

0.5

I .0

I .5

2.0

2.5

3,0

3.5

4.0

4.5

5.0

C female "whiny" calls

5
4

D female "whiny" calls

3 2
1

0.5

I .0

I .5

2.0

2.5

3.0

3.5

4.0

4.5

5.0

654

male "guttural" call

F female rattle call

................................................

G rattle call

3 2
1

0.5

I .0

I .5

2.0

2.5

3.0

3.5

4.0

4.5

5.0

Time

(seconds)

FIC. 5. Vocalizationsof Suiriri s. affinisfrom varioussitesin Amap (AP) and Mato Grosso(MT), Brazil. MPEG numbers parentheses in matchtaperecordings with specimens the MuseuParaense in EmilioGoeldi, Be16m, Par, Brazil. (A) Male (MPEG no. 54881)loudsong(MT, near Cuiab, 22 September 1999).(B) Female loudsong(MT, near Cuiab, 21 September 1998).(C) Female"whiny" calls(MT, near Cuiab, 22 September 1999). (D) Female"whiny" calls (AP, north of MacapP,12 February1999). (E) Male guttural call (MT,

January 2001] 10 Suiriri suiriri finis qf.

A male/female duet
...

NewFlycatcher Cerrado from Regwn

69

male/female loudsongs

female"whiny"calls

0
10
8 6

1.0

2. 0

3;0

4.0

5.0

6JO

7:0

8' .0

9' .0

10.0

Sui suid affinb

male/female duet

male/female loudsongs

female "whiny" , } calls

....
1.0

2.0

3.0

4.0

5.0

6.

i0

7.0

8.0

9.0

10.0

Suiriri suiriri afJnis

male/female duet male/female loudsongs

female 'lwhiny" calls

1.0

2.0

3.0

4.0

5.0

6.0

7.0

8.0

9.0

10.0

10 Suiriri suiriri bahiae

D male/femaleduet
female "whiny" call female loudsong maleloudsong

I .0

2.0

3.0

4.0

5.0

6.0

7.0

8.0

9.0

lO.O

Time

(seconds)

FIG.6. Male-femaleduetsof Suiriris. affinis and S.s. bahiae from various localities Amap (AP), Mato in Grosso (MT), and Pernambuco Brazil.(A) S.s.affinis (PE) from Chapada dosGuimarfies, (21 September MT 1999).(B) S.s.affinis from Chapada dosGuimarfies, (21 September MT 1999).(C) S.s.affinis from nearPorto Grande,AP (7 February1999). (D) S.s. bahiae from LagoaGrande,PE (15 January1997).All recordings by
Kevin J. Zimmer.

displaying.Duets almostalwayswere initiated by severalnasal,single-notefemale calls (Fig. 6A-D) given in rapid succession. the male If

male,land nextto her, and thenthe pair would simultaneouslysing their respective loudsongs. The postureof both birdsduring duets wasnearlyhorizontalwith thewingstypically was in a differenttree or anotherpart of the drooped slightly at the sides.Birds usually sametree,he would rapidly fly toward the fe- quiveredtheir wings slightly while keeping

S.s. bahiae exhibited

identical

behaviors

when

Chapada dosGuimarfies, September 21 1999).(F) Rattlecall from female(MT, nearCuiab, 22 September 1999).(G) Rattlecall of bird of unknownsex(AP, north of Macap& 12 February1999).All recordings by
Kevin J. Zimmer

70

ZIMMER, WHITTAKER, OREN AND

Suiriri suiriri bahiae

[Auk, Vol. 118

A male loudsong
4-

0 o lO 987-

0.5

1.

1.

2.0

2.5

3.0

315

410

4. 5

B female loudsong

65
4

3
2

1 0

0.5 I

1.0

1.5

'1

2.0

2.5

3.0

3.5

4.0

4,5

5.0

C male "guttural" call

D male "pitty-chew"

0.5

'

1.

'0

I .5

'

2.0

'

2.

3.

'0

3.5

'

4.0

'

4.5

'

5.0

7-

E female "whiny" calls

654-

3
2

1 0

0.5

1.

1.5

2.

2.5

3.0

3.5

4.0

4 5

5.0

Time

(seconds)

FIC. 7. Vocalizations Suiriri s. bahiae of recordednear LagoaGrande,Pernambuco, Brazil. (A) Male loudsong(25 January1999).(B) Femaleloudsong(25 January1999).(C) Male guttural call (25 January1999).(D) Male "pitty-chew" calls(25January 1999).(E) female "whiny" calls(25January 1999). recordings Kevin All by
J. Zimmer

January 2001]

NewFlycatcher Cerrado from Region

Suiriri suiriri suiriri

71

7-

A presumed loudsong
.....

65 4 3 2 1 0

.....
0.5

'" ................... .............

i
1.5

1.0

2.0

2.

15

3.0

3.5

4.0

4.5

5.0

B single-note call

C single-note call

D single-note call

E single-note call
.;;'. -...

... j'..'.
I i
I

0.5

1 0

1.5

2 0

2.5

F rattle call

.....................

0.5

1.0

1.5

2.0 Time

2.5 (seconds)

3.0

3.5

4.0

4.5

5.0

FIG.8. Vocalizations Suiriris.suiriri.(A) loudsong of (Argentina, Salta, October 6 1987; Hilty recordS. ing). (B)Single-note (Argentina, call Salta,6 October 1987; Hilty recording). Single-note (Brazil, S. (C) call

RioGrande Sul,50kmsouthwest Antonio Miss6es,November W.Belton do Santo das 4 1975; recording, LNS
19531).(D) Single-note (Bolivia,depto.SantaCruz, westof SanIsidro,6 October1983;T. Parkerrecordcall ing, LNS 33629). Single-note (Paraguay, (E) call Chaco Province, Filadelfia, August1990; Finch 21 D. record-

ing,LNS57891). Rattle (Paraguay, (F) call Chaco Province, Filadelfia, August 21 1990; Finch D. recording,
LNS 57891).

themdrooped, sometimes and simultaneouslybackby flyingin andperching nearus without shivered tail up-and-down a shallow the in arc vocalizing. Even after leaving and being of less than5.UnlikeS.isIerorum, neither affinis broughtbackby tape playbacktwo or three norbahiae habitually selected prominent perch- times, those males wouldnotsingontheirown. es from which to display,and often duetted In each case, female a eventually in giving flew fromwithinthecanopy trees. of Birdsrespond- several nasalcallsand the pairs immediately ing to tape playbackoften sangas they were beganto duet.
flying towardthe soundsource, behavior a that
we did not observe in S. isIerorum. On several

Tape playback experiments.--Playback experi-

ments with Suiririrecordings offerfurtherevoccasions had affinis we malesthatweresepa- idenceof the significance vocal differences of rated from their matesrespond tape play- among taxa,and of their role aspotential to the

72

ZIMMER, WHITTAKER, OREN AND

[Auk,Vol. 118

TABLE Summaryof playback 3. trials (asdetailedin the text) involvingSuiririflycatchers islerorurn, (S. S.s. affinis, bahiae S.s.suiriri).Numbers parentheses S.s. and in indicate number pairsor familygroups the of of the subject taxonthat were testedfor eachstimulustaxon.
Response playbackof to

Locality

islerorum

affinis
S. islerorum

bahiae

suiriri

Mato Grosso, Brazil Amapa, Brazil Mato Grosso, Brazil Totals Pernambuco, Brazil

strong(11) none(3) none(12) none(15) none(5)

none (11)
S.s. affinis strong(3) strong(12) strong(15)
S.s. bahiae

none(11) strong(3) strong(12) strong(15) strong(5)

none(6) none(2), moderate(1) -none(2), moderate(1) none(3), strong(2)

strong(5)

isolating mechanisms. We performed tape to whichnoneof themresponded. thenpreWe playbackexperiments 11 pairs of Suiriri is- sentedthem with playbackof nominateS.s. on lerorum near Cuiaba and ChapadadosGuimar- suiririfrom Argentina.The adult pair from the aes,Mato Grosso,Brazil (six pairs in Septem- first family group responded approaching by ber 1998and five pairsin September 1999).The the sound source peeringabouton two ocand six pairsin the 1998trialswerepresented with casions, but did not vocalize. None of the four recordings S.s.bahiae of from Pernambuco, Bra- birdsfrom the otherfamily groupshowed any
zil, S.s. affinisfrom Cuiab, Mato Grosso,Brazil and S.s. suiriri from Argentina. The five pairsin the1999trialswerepresented with the samerecordings S.s. bahiae S.s. affinis. of and None of the 11 pairs tested showedany response tapeplayback suiriri,affinis, bahto of or iae vocalizations. When presentedwith playback of prerecordedS. islerorum vocalizations, all 11 pairs responded by immediately apinterestin playbackof nominateS.s.suiriri, nor

did thelonepair.Whenpresented with tapeof

S.s. affinisfrom Mato Grossoand S.s. bahiae from Pernambuco, threepairsof adult S.s. all affinis from Amapa responded stronglyby approaching and duetting. In January 1999,we performedtapeplayback

experiments five pairs of S.s. bahiae on near Lagoa Grande,Pernambuco, Brazil. Thosebirds proaching sound the source duetting and mul- were first presentedwith taped vocalizations tiple times.In severalcases, thosebirds flew in of S. islerorum from Mato Grosso, Brazil. None from >100 m away when responding play- of thefive pairsshowed response those to any to
back of their own vocalizations.

playbacks. thenpresented We themwith play-

back of S.s. affinisfrom Mato Grosso.All five pairsresponded stronglyto playback S.s.afof iabaand ChapadadosGuimaraes, Mato Gros- finis,alwaysapproaching sound the source and so,Brazil (six pairs in September 1998and four duetting the fourthplayback. alsopreby We pairs and two family groupsof four in Septem- sented fivepairsof S.s.bahiae the with playback ber 1999).Birds in all 12 trials first were pre- of nominateS.s. suiriri from Argentina.Three sentedwith taped vocalizations S. islerorum of the pairs did not respondto the voice of of from Chapada dos Guimaraes, then with re- nominate birds.The othertwo pairsresponded cordingsof S.s. bahiae from Pernambuco, Bra- strongly (approachand vocalizations), but it zil. Noneof the 12 pairs or family groups re- shouldbe pointedout that theywereprevioussponded to playbacksof vocalizationsof S. ly stimulated tapeof affinis by from Mato Grosislerorum, all 12 pairs of adultsresponded so.Theresultsof those but playback trialsaresumstrongly to playback of S.s. bahiae vocaliza- marized in Table 3. tions.We alsoperformedtape playbackexperForaging behavior social and structure.--Suiriri iementson two family groupsand one pair of islerorum typically was encountered closely in S.s. affinisnorth of MacapP, Amap, Brazil associated pairs,apartfrom otherspecies. They (February1999). Those birds first were pre- foragedmostlyin trees(rarely descending to

We performedtape playbackexperiments on 12 pairs or family groupsof S.s.affinis near Cu-

sented with playback S.islerorum of recordings, low shrubs the ground),movingenergetior

January 2001]

NewFlycatcher Cerrado from Region

73

cally with shortpauses betweenhopsand frequent and erratic changes direction.They of gleaned various smallarthropod preyfrom leaf and branchsurfaces. Once prey was spotted, they frequently used wing-assisted hops to close distance rapidly,thenlungedforwardfor a horizontal capture,or jumped upward to gleana preyitem from overhead. Theyalsoengagedin frequentdarting aerialsalliesupward or outward to glean prey from undersides of
leaves or branches. Those sallies varied from 15

ilar to foragingtechniques routinelyemployed by bluebirds(Sialiaspp.;K. Zimmer pers.obs.) in North America.Suiriris.affinis alsomadeoccasionalshort, looping sallies of 1-2 m out from crowns of trees in pursuit of flying insects, that behavior but waslessfrequentthan in S. islerorum. Like S. islerorum, affinis periodically wagged its tail downward in a relaxed mannerwhenforaging,particularlyafterperch changes. Foragingbehaviorof S.s. affinisobservedin Amap, Brazil was identicalto that
described above from Mato Grosso.

cmto 1 m and typicallyinvolvedlittle truehovering.Thosebehaviors were similar to that of membersof the generaTolmomyias Platyrand inchus, althoughS. islerorum were more constantlyin motion and were more agile and acrobatic in pursuing prey (K. Zimmer pers.

The socialstructureof S. islerorum requires further investigation. have never encounWe tered more than two birds together.However, in September1999, we had two experiences that suggestthat S. islerorum at least occais observ.).Suiriri islerorum also made occasional sionally polygynous. On 21 September1999, alongthe Agua longer(up to 2 m), loopingsallies from the we taped a pair of S. islerorum out crown or lateral branches of trees and back to Fria Road,Mato Grosso, Brazil. After obtaining the pursue flying insects. Perch changes wereusu- vouchertapes,we collected displayingfeally followedby an obvious,but relaxeddown- male (MPEG 54867).The male flew a shortdisward dip of the tail, similar to tail movements tanceout of sight.Immediately afterretrieving more tape playback of Vermilion Flycatcher (Pyrocephalus rubinus). the female,we broadcast Suiriris. affinis the sameareasfrequently from the samespot, and a male returned.As in perch-gleaned hover-gleaned or small fruits soonas the male perchedin front of us, a sec(particularlythoseof Curatella americana), al- ondbird flewintothesame treeandsanga typthoughthey spentmore time in pursuit of ar- ical female loudsong.The male immediately thropods. Suiriris.affinis foragedmostlyin rel- chased that second female, pursuing her This wasrepeated on ativelyopentrees,progressing hopsof 5 to aroundtreesand shrubs. by each time we presentedtape 15 cm, primarily alongmain branches. Their five occasions; movements throughtreesweregenerally more playbackof duets the male would approach, by female, whichhe leisurelyin pace (almost Vireo-like)and more followedclosely thesecond drive off. Only once predictable progression in than in S. islerorum. would then aggressively Birds pausedfor 2-15 s betweenhopsto scan during those encounters did the two birds live foliage, fruit clusters,or bark surfaces. duet. On our last playbackattempt, the male They generallyworkedfrom the interiorof a drove the femaleto the groundand appeared tree outward, or from the subcanopy but to mounther.They then flew to a nearbytree up, withoutfrequentchanges directionas in S. and performedmore duetsbeforewe collected of islerorum. Arthropodpreyusually(-75% of all both birds (male MPEG 54869, secondfemale captures)were gleaned directly from leaf or MPEG 54868). Gonadsof all three birds were branchsurfaces reachingout, up, or down. enlarged. by Later on 21 September, were collecting we Foraging birds oftenpickeddirectlyat branch surfaces, particularly on trees with furrowed farther along the Agua Fria Road when we Once bark. Theyalsofrequently hover-gleaned prey taped in another pair of S. islerorum. by hovering1-2 s beneathgreenleaves the again,we collected female(MPEG 54872) in the outerportionof the canopy. Thatwasdifferent first, and the male flew off. After a few minutes from the darting sallies(with little or no true we presentedmore playbackof duet vocalizaflew hovering)madeby S. islerorum. Suiriris. affinis tions. Immediately,a pair of S. islerorum often dropped 1-3 m from branchesto the into the same tree from which we had collected groundto pounceon prey spottedfrom above. the female, and proceeded to duet multiple That behaviorwas particularlycommon re- times. After obtainingvouchertapes,we colin centlyburnedor very openareas,and wassim- lectedboth birds (secondfemaleMPEG 54873,

74

ZIMMER, WHITTAKER, OREN AND

[Auk, Vol. 118

male MPEG 54874).The first femalehad a con- fying the sexof a territorialtresspasser (Faraspicuous brood patch and the gonadsof all baugh1982).A likely byproductof sexualspecthreebirds were enlarged. ificity of duet roles is the strengthening of In each of those instances, what we assumed reproductiveisolation.In an habitually duetto be the originalmaleappeared returnwith ting genus to such Suiriri,vocaldifferences as bea secondadult female in breeding condition tween taxawould seemto presentmajorobstawithin 5 min of the time that we had collected cles to reproductive compatability.Further his previous mate,which suggests those barrierswould be presented significant that by difmaleshad beenmatedto two birdsall along.It ferences stereotyped in physicaldisplaysas-

seems highlyunlikelythatmorethan onemale

was involved in either case. There was no evi-

sociated

with

the vocalizations.

As detailed

above,pairs of S. islerorum perform distinctive dence a second of territorialmaleor pair in the displays with eachduet.Nothingthat we have vicinity prior to collection the first females. heard or seenin the vocal and display reperof Had we been at a boundary between two ter- toire of affinis bahiae or suggests approach an to ritories,it seems likely that our tape playback islerorum. wouldhaveelicitedresponses frombothneighThe biologicaland taxonomic significance of boringpairs.Instead,playback broughtan im- thosedifferences suggested the resultsof is by mediate approachand duet response from a our playbackexperiments. Althoughwe were singlepair. Followingour collection the fe- unableto perform all pairwisetests,the ones of male of eachpair, the respective malesflew off performed wereunequivocal. Noneof the affinout of sight,after which a male flew backfrom is or bahiae testedshowedany response reto roughly the same area minutes later in re- peated playbacksof islerorum, did any of nor sponse further playback.Birds responding the islerorum to respond playbacks the other to of

to the second roundof tape playback perched in the same adjacent or shrubs those as usedby the birds respondingto the first tape playbacks. haveonly rarely foundpairs of S.isWe lerorum occurring close in enough proximityto oneanother thatwe couldhear(evendistantly) two pairs from a single spot, making it even

taxa.

Morphologicaldifferences from affinisand bahiae are subtle yet distinct, even when observed the field.Amongthe eightcharacters in
that were measured, Suiriri islerorumdiffered

significantly from affinis culmenlength,bill in width, bill depth, wing chord, tail length, less likely that two independent territorial width of the central rectrix, and width of the pairswereinvolved eachinstance. in (Table2). The two speSystematic relationships.--Some generaliza- pale tip to the rectrices ciesalsodifferedin the colorpatternof the outtions regardingrelationships amongthe yeler rectrices, and in whether or not the central low-belliedgroupof Suiriricanbe made.Suiriri rectrices werepale-edged. Our sample affinis of islerorum sufficientlydistinctto be considis indicated substantial sexualdimorphism bill in ereda separate species underanyof the widely size and color of the mandible, whereas no such accepted species concepts (McKitrick and Zink wasapparentin islerorum. is not It 1988). Vocal distinctions are particularly dimorphism share strong.Suiriri islerorum differsstrikinglyfrom known whetherjuvenilesof S. islerorum nominatesuiriri, affinis,and bahiae its male the unique spottedpattern to the upperparts in and femaleloudsongs, duets,and in all known foundin juvenilesof otherSuiriri. Most important,Suiriri islerorum S.s. afand male and female calls. Duets serve multiple finis are syntopic multiplesites,andshowno at functions, including coordinateddefenseof of At territory and pair-bonding. Suiriri, asin the evidence interbreeding. the two localities In majorityof duetting species, same the typesof in Mato Grosso, Brazil, where we conducted duets are used during boutsof territorial de- field studies,both forms were relatively comfense,as a prelude to copulation,and when mon and all pairs of Suiriri were assortatively type pairs reunite followingvisual separation (Far- mated by vocaltype and morphological abaugh1982,K. Zimmer pers.observ.). sex- (as confirmedby tape recordings,specimens, In Thus, they meet the priually monochromatic species (Suiriri), sexual and tissuesamples). species unspecificity of duet contributions probably mary criteria for being considered serves in mate attraction as well as in identider the biological species concept.

January 2001]

NewFlycatcher Cerrado from Region

75

Finally, preliminary molecular analyses of tissue samples indicates that S. islerorumis stronglydifferentiated genetically from both S. s. affinisand S.s. bahiae Batespers. comm.). (J. Those results will be presentedelsewhereas part of an overview of genetic diversity in
Suiriri.

Basedon vocal similarities,S.s. affinisand S. s. bahiae clearly are closelyrelated. The latter formresembles formerin havinga longbill the and yellow belly, but differs in being shorterwinged, showing little back-rump contrast, and in lackingthe contrasting palebaseto the rectrices (Traylor 1982). However, our examination of 61 specimens affinissuggests of that the degreeof back-rumpcontrast and the extent of the contrasting pale baseof therectrices are individually variable characters, and extremeexamples affinis of may at leastapproach typical examplesof bahiae. Hayes (2001) also found significantplumagevariability in specimens of bahiae. vocalizationsof bahiae All appear to be homologous thoseof affinis, to and displaybehaviors duettingpairsare identiof cal. Furthermore,in reciprocal tape playback experiments, all pairs of bahiaeresponded stronglyto playbackof affinis vocalizations, as did all pairs of affinis vocalizations bahiae. to of The relationshipof nominateS.s. suiriri to the yellow-bellied forms remains uncertain. Nominate birds are strongly differentiated morphologicallyand vocally from islerorum, and in tape playbackexperiments, latter the species did not respondto playbackof suiriri
vocalizations. There is no reason to consider

responded to suiriri vocalizations,whereas most pairs showedno response. More comprehensiveinventoriesof tape recordedvocalizationsof nominatebirds are neededfor a proper vocal analysis. Also needed are field studies from the contact zonebetween suiririand affinis, as well as more extensiveplaybackexperimentsfocusedon this pair of taxa. Molecular analysesin progress (J. Bates et al. unpubl. data) alsoshouldclarify thoserelationships. Ecology conservation.--Suiriri and islerorum is a bird of the cerradoregion, a biogeographic region that encompasses to 2 million km2 1.5
centered on the Brazilian shield, with extensions into eastern Bolivia and northeastern Par-

aguay (Eiten 1972). The term "cerrado" has been applied generallyto virtually all of the nonforest habitats thatregion,although of geographicallyit variesgreatlyin species composition and physiognomy. Five major subtypes of cerradohave been recognizedby botanists (Eiten 1972).Of those,we havefound Suiriri islerorum occurin three: (1) cerradosensu to stricto, woodland with closed scrub and scattered

them as anythingotherthan separate species.

In the absence of contact, nominate birds also

would appearto be sufficiently distinctfrom affinis and bahiae bothmorphological vocal in and characters to warrant separatespecies as status. However,the statusof intergrades hyor
brids from the contact zone in northern Para-

trees;(2) campocerrado, moreopenscrubwith a few trees;and (3) camposujo,grassland with scattered shrubsand few trees.In the Chapada dos Guimares and Cuiab regionsof Mato Grosso,Suiriri islerorum locally fairly comis mon in someareasbut seeminglyabsentfrom others.Moderatelyclosed,shrubbyareaswith a significant grass component and scattered trees2-5 m tall (asexemplified the first sevby eral kilometers theAgua Fria roadnearChaof pada dos Guimares)have the highestdensities of S. islerorum. Our limited surveyshave averaged to 2 pairs per kilometerof road, but 1 more extensive dawn surveys using systematic tapeplayback undoubtedly wouldyield higher densities. The species somewhatless comis mon along the Coxipo do Ouro road near Cuiab, where the habitat is more wooded with

guay and southwesternBrazil remains an issue.Hayes (2001) found that presumedhybridsfrom the contact zonehad plumagecharactersintermediatebetweensuiriri and affinis, and exhibitedsignificant plumagevariability. The apparentscarcityof parentalphenotypes from within the contactzone suggested that the two forms were freely interbreeding, and therefore, conspecific (Hayes2001).Furtheruncertaintyis introducedby our playbackexperiments, whichsome in pairsof affinis bahiae and

pockets regularlyspaced of trees(closer certo rado sensu stricto;Eiten 1972). Suiriri s. affinis occursalongside islerorum both areas,but is in more common in the more open woodlands and less commonin the shrubbier,grassier
cerrado.

The areas in which we have found Suiriri is-

lerorum subjectto regular human perturare bation.The biggest disturbance duringthetenure of our work has been fire. Accidental or

deliberatelyset fires have burned significant

76

ZIMMER,WHITTAKER, AND OREN

[Auk, Vol. 118

portions cerrado of along AguaFriaroadin the

three of the last five years, and have also affectedthe nearbynationalpark. Localpopulations of both Suiriri islerorum and S.s. affinis havepersisted heavilyburnedareas, the in but repeatedburning of the sameareaswithout sufficienttime for recoveryof the shrub community likely will havea negative effect.Hous-

incompleteknowlege of distributionsmay result in the serious underestimation regional of and local biodiversity and endemism.Such miscalculations could have seriousrepercussionswhen important conservation priorities

are beingset.In an analysis the intensityof of collectingefforts in the cerradoregion,Silva (1995)found that roughly70% of the cerrado ing developments spread have into the cerrado regiondid not meetthe criteriaof havingbeen along the Agua Fria road, and continued ex- "minimally sampled"for birds.It is instructive pansionsoonwill threatenthe type localityof that Suiriri islerorum occurs in at least seven of islerorum. the sitesthatmet Silva's requirements being for sampled,including three reOn a broaderscale, dry forestand grass- satisfactorally the land habitats of central South America are congions (Goiania,Cuiab, and ParqueNacional sidered both amongthe mostendangered and Noel KempffMercado)with a high densityof localities, yet, hasesand unique biomesin the Neotropics(Stotz et al. minimallysampled by 1996). The cerrado is especiallythreatened, caped detection field workers.If a cryptic with more than 75% of the 41 endemic birds species bird could be missedin the most of considered at risk, and more than 45% consid- heavily sampled30% of the cerradoregion, biodiversity ered eitherthreatened endangered or (Stotzet then the potentialfor overlooked areas mustbe al. 1996).The primary threatderivesfrom the in the remainingunder-sampled wholesale mechanized conversion of native great. Our recognitionof Suiriri islerorum a as crypticspecies initially basedon an apprewas cerradoand campocommunities large-scale to agriculture,particularly soybeans. The dry- ciation of vocal differences within Suiriri over range.It is criticalthatfield season practice annuallyburningopencoun- a broadgeographic of conducting surveys the cerrado of try habitatsfor small-scale agriculture,deeply personnel and other endangered habitats have access to ingrained amongmost rural populations,is a adequate tape recording equipment and be secondary threatwhoselongtermenvironmenof or tal consequences have only recently raised proficientin the recognition bird voices,
widespread concern.
Before the true conservation status of Suiriri

more taxa inevitably will be undersampledor overlooked (Parker 1991).

ACKNOWLEDGMENTS

islerorum can be assessed,we first need a more

complete understanding its distribution. of We expect that it will eventually proveto bebroadly sympatric with S.s. affinis throughout much of the cerradoregion.At present,Suiriri islerorum is known to occur in two large national parks,ChapadadosGuimaraes NationalPark, Mato Grosso, Brazil, and Noel KempffMercado NationalPark,depto.SantaCruz,Bolivia.Both parks contain extensive areas of seemingly suitablecerradohabitat,and shouldbe systematically surveyedto assess densitiesof Suiriri
islerorum and other cerrado endemics. Another

We are particularly grateful to Dionisio Pimentel (MPEG), who skillfully handled the collectingand specimen-prepping duties on our collecting trip. T. Schulenbergwas a constant source of advice throughout project,and alsoprovidedassistance this in obtaining somekey references. Batesalso proJ.
vided much useful advice, and facilitated the iden-

protectedarea that shouldbe surveyed isfor lerorum the ReservaEco16gica is Serra das Araras in western Mato Grosso, Brazil.

tification of plants from our study areas. Special thanksmust go to M. and E Isler, who generously donatedtheir time and talentsto providingthe map and spectrograms (respectively) usedin this paper K. Garrett (LACM) provided criticalassistance coin ordinatingspecimen loansfrom several institutions.
We also thank J. V. Remsenand S. Cardiff (LSUMZ);

T. Schulenberg, Willard, and S. Hackett(FMNH); D.

E C. Sibley(YPM); C. Cicero(MVZ); and R. Ridgely and D. Agro (ANSP) for the loan of specimens from their respectiveinstitutions.A. Schaffnerassisted with the statistical analysis morphological of data.G. Budney, Bridgeford, Priori, and the restof the T. A. staff at the Library of Natural Sounds, CornellUni-

The need for adequate surveys of endan- R. K. Panza (CM); S. L. Olson and P.Angle (USNM);

geredcerrado habitats beenpointedoutby has

others (Bateset al. 1992, Silva 1995, Parker and Willis 1997),but its importance cannotbe overstated.Lack of understanding species of limits

in widespread, polytypicgroups,alongwith

January 2001]

NewFlycatcher,from Cerrado Region

77

versitywere of great help in allowing access the to los (Rhinocryptidae), with descriptions the of LNS collection recordings. of Thanksalsoto S.Hilty, Ecuadoriantaxa, including three new species. R. Ridgely,and M. Cohn-Haftfor sharingadditional Pages47-88 in Studiesin NeotropicalOrnitholrecordings with us. D. Lane painted the attractive ogy Honoring Ted Parker(J.V. Remsen,Jr.,Ed.). colorplate.B. Carlosprovided transportation our on Ornithological Monographs 48. no. collecting trip. We thank E. Goodale,D. Kroodsma, KROODSMA, E. 1984.Songs the Alder Flycatcher D. of T. Schulenberg, an anonymous and reviewerfor their (Empidonax alnorum) Willow Flycatcher and (Emhelpful commentson earlier drafts of this manupidonax traillii) are innate.Auk 101:13-24. script.Finally,we gratefullyacknowledge Eman- KROODSMA, E. 1986.Designof songplaybackexV. D. uel for providing us with travel opportunities that periments. Auk 103:640-642. havemademuchof our work possible. KROODSMA, E. 1989.Male EasternPhoebes D. (Sayornis phoebe, Tyrannidae, Passeriformes) fail to LITERATURE CITED imitate songs. Journalof Comparative Psychology 103:227-232.
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