NielsKrabbe 219 Bull. B.O.C. 2008 128(4) Vocal evidence for restitution of species rank to a Santa Marta endemic: Automolus rufipectus Bangs (Furnariidae), with comments on its generic affinities by Niels Krabbe Received27November2007 Some passerines in the family Tyrannidae possess entirely innate songs (Stein 1963, Lanyon 1978, Kroodsma 1984, 1989, Kroodsma & Konishi 1991), with no local dialects. A similarlackofdialectsinseveralotherfamiliesstronglysuggeststhatsongisinnateinnear- ly all suboscines, which has led to a recent re-evaluation of the taxonomic ranks of many forms (see, e.g., Remsen 2005). Several of these are members of the Furnariidae (Vaurie & Schwartz 1972, Whitney & Pacheco 1994, Kratter & Parker 1997, Zimmer 1997, 2002, Robbins & Zimmer 2005, Zimmer et al. 2008), which includes a large number of taxa that occur in dense and dark habitats, where vocalisations, rather than visual signals, probably play a major role in species recognition. The furnariid taxa Furnarius cinnamomeus, F. lon- girostris, Synallaxis maranonica andHyloctistes virgatus are afforded speciesrankonthebasis ofvocalcharactersbysome authors,whereas others (e.g. Remsenetal. 2007) awaitthepub- lication ofvocal studiesbefore accepting taxonomic changes. DuringfieldworktodocumentthevoicesofbirdsintheSierraNevadadeSantaMarta, northern Colombia, in February 2007, 1 ascertamed that the songs of several endemic taxa presently treated as subspecies are quite distinct from purported conspecifics. The present paperaddressesAutomolus rubiginosus rufipectus,describedasafullspecies.RuddyFoliage- gleaner A. rufipectus, by Bangs (1898), which differs so much from its alleged conspecifics that even its generic placement may demand re-evaluation. A. rufipectus was ranked as a subspecies ofA. rubiginosus by Hellmayr {in Cory & Hellmayr 1925), a treatment that has beenwidely followed until the present. Methods m I tape-recorded songs of four individuals of rufipectus at 1,200-1,800 in the recently established ProAves Colombia reserve 'El Dorado' on the San Lorenzo ridge in the north- westernpartoftheSierraNevadadeSantaMarta,dpto. Magdalena,Colombia(11°06-08'N, 74°03-06'W). Birdswerefirsttapedundernaturalconditionsandthenattractedusingplay- back for identification. Three of the four recorded birds were seen, and one was photographed (Fig. 1)nearitsnest. Eightmonthslater,anindividualwascollectednearthis nest (C. D. Cadena, specimen in Universidad de Los Andes, Bogota, catalogue number ANDES-O-0433). RecordingsweremadeusingaSennheisserME67microphoneand aSony TCM5000 tape recorderwith Type I cassettes. Two additional recordings of rufipectus were consulted (Boesman 1999, Strewe et al. 2003). Sonograms were compared among rufipectus and all other species or vocally distinct subspecies orsubspecies-groups ofAutomolus,Hylocryptus,Anabazenops and Syndactyla, the latter three genera all encompassing species formerly included in Automolus. Sonograms were generated using the sound-editing program CoolEdit Pro (Syntrillium Software), whichwas also used for vocalisation analysis. Observations onhabitatandbehaviour were noted in the field and are compared with published information. NielsKrabbe 220 Bull. B.O.C.2008128(4) Results — Vocalisations. Atotalof196songphrases(from5-6differentindividuals)ofrufipectuswere analysed. Songs consisted of resonant notes (3-4 harmonics), given in a series of4-6 simi- lar notes, loudest at 1,800-3,000 Hz (second or third harmonic). Individual notes were up-downstrokes, the downstroke slightlymorepronounced, and thepeakvolume athigh- est pitch. The loudest pitch was the second harmonic (1,800-2,000 Hz) in two or three individuals, the third harmonic (2,400-3,000 Hz) in two individuals, and second and third equally loud in one individual. The song was usually given in sets of two (less commonly one, rarely three) phrases, with a brief pause of 0.3-0.5 seconds between the two phrases. Thefirstofthetwophrasesfrequentlyhadanextra,oftenslightlyweakerandlowerpitched introductory note, and was thus longer (0.42 ± 0.04 seconds) than the second (0.36 ± 0.03 seconds).Thedoublephrasesweredeliveredfairlyregularlywithpausesbetweenthem2-4 seconds long. All the individuals of rufipectus compared had similar songs, differing in virtually all aspects from songs ofother forms ofAutomolus rubiginosus (Figs. 2a-i, 3a-b), the only simi- larities being the length of the song phrases and the tendency of rufipectus to deliver two songphrasesinsuccession,atendencyalsonoticedinsomeindividualsofotherformsofA. rubiginosus from both Central America {rubiginosus, veraepacis, saturatus) and the Amazon (unnamed taxon). The song of rufipectus was more similar, indeed remarkably so, to the song ofHylocryptus erythrocephalus (Fig. 3b-c). Calls of rufipectus recorded were churrs, two- to three-noted whilst foraging (Fig. 4d) and three- to four-noted when given in alarm (Fig. 4e), at 0.8-2.0 kHz (first and second, or second and third harmonic), lower pitched than song and differing distinctly from calls of bothA. rubiginosus (Fig. 4a-b) and Hylocryptus erythrocephalus (Fig. 4c), which in turn have fairly similar calls except for the latter being lower pitched. No call resembling the high- pitched (4 kHz) call (not show^n) purportedly given by H. erythrocephalus (Coopmans et al. 2004, cut5; misidentified?) was ever heard from rufipectus. Figure1.Automolusrufipectus,SierraNevadadeSantaMarta,Colombia,February2007(JoeTobias) NielsKrabbe 221 Bull. B.O.C.2008 128(4) Songsofeightoftheother 13currentlyrecognisedsubspeciesofA. ruhiginosus (Remsen 2003), aswell as ofanunnamed taxonfrom the Amazonianlowlands, were compared (Fig. 2a-i). They all consist of a single, 0.3-0.6 seconds-long, distinctly rising note, sometimes (withintra-individualvariation)possessingashortintroductorynote. Asinotherspeciesof Automolus, the notes are highly resonant (with 3-4 audible harmonics), and as in rufipectus the pitchis variable. The single recording ofthe unnamed taxon from the Amazonian low- lands (all other forms being lower montane) is decidedly higher pitched (Fig. 2i), thus suggesting an isolated position of this form. The five forms for which no recordings were avilable at the time of the study wereguerrensis,fumosiis, cinnamomeigula, venezuelanus and caquetae. Songs ofthe former four are described asbeing similar to songs ofadjacentforms (Howell & Webb 1995, Stiles & Skutch 1989, Hilty 2003; A. M. Cuervo pers. comm.). Only caquetae appears to be vocally unknown, but its poor morphological differentiation from, and seeminglycontinuous distributionwith brunnescens suggestthatitmaybealso similar. — Natural history. On three occasions rufipectus was observed foraging. It remained within m 0.5 ofthe ground, and spentmostofitsforagingtime onthe ground, rummagingamong large dead leaves as does Hylocryptus erythrocephalus (Ridgely & Greenfield 2001, Remsen 2003;pers. obs.),onlyoncepeckingbrieflyatadecayingtrunkasdoesAutomolusruhiginosus regularly (Remsen 2003; A. Tobias pers. comm.). This concurs with Todd & Carriker J. (1922), who wrote that it 'does little climbing, but keeps hopping about on or near the ground". A verticalbank with several entry holes 1-2 m above was found. Thebird would always leave from the same hole when disturbed, but when entering, it would usually briefly enter one ofthe other holesbefore moving to the final one. — Habitat. A. rufipectus was found in forest and second-growth habitats, which were transitional between the dryAcacia scrub at sea level and the humid forest above 2,100 m, but more similar to the latter. The undergrowth was dense, and the litter included many m largeleaves. InFebruary,adrymonthwithoccasionalrain,thehabitatat1,200-1,800 was reminiscent of dry-forest haunts of Hylocryptus erythrocephalus in south-west Ecuador and north-west Peru. On two occasions I found rufipectus in heavily disturbed areas. Todd & Carriker (1922) noted: Tt is partial to the heavy forest or thick second growth, and is fond m ofdarktanglesandravines'. Ididnotfinditoritshabitatabove 1,800 ontheSanLorenzo ridge 70 km east-north-east of Pueblo Viejo (the type locality). This is evidently in agreementwithCarriker's experience; although the type ofrufipectus was originallysaid to have been taken at c.8,000 feet [2,438 m] (Bangs 1898), Todd & Carriker (1922) gave the species' altitudinal range as 3,000-6,000 feet [914-1,829 m], and added the note '7500 feet, according to Mr Brown' [the collector of the type]. The nest I found on the San Lorenzo ridgewas atc.1,750 m. — Distribution, population and conservation. All records are from the north-western and northern slopes of the Santa Marta Mountains. Collecting sites are from around the San Lorenzoridge (Cincinnati;LasVegas;AguaDulce) andintheupperrfoAnchadrainage (La Concepcion; San Antonio; Pueblo Viejo; Chirua; Heights of Chirua) in dptos. Magdalena andGuajira (Todd &Carriker 1922). These and some additional, more recentsightorvocal j records were mapped by R. Strewe, G. Lobaton & S. Sanchez, of Fundacion ProSierra Nevada, in 2004 (see http://www.alpec.org/Atlas/Tama%F1oFinal/Automolus I rubiginosus.jpg). However,thesouthernslopesofthemountains (largepartsofwhichhave j been converted for coffee farming) have been subject to very little ornithological j investigation to date. I found the species widespread but uncommon on the San Lorenzo i 1 NielsKrabbe 222 Bull. B.O.C.2008 128(4) ^ 5 Z 3 UDl O 2 LU ill it CC ^'^"^ li- 1 \ i i ^ 10 13 TIME(sec.) Figure 2. Songs of 12 taxa of foliage-gleaners. A-J ten different subspecies of Automolus rubiginosus: A: rubiginosus (Oaxaca,Mexico;P. Boesman). B: veraepacis (Chiapas,Mexico;P. Boesman). C: saturatus (Darien, Panama;L. R. Macaulay). D: nigricauda (Pichincha,Ecuador;N. Athanas). E:sasaimae(Boyaca,Colombia;O. Laverde). F: obscurus (Surinam; O. Ottema). G: bninnescens (Zamora-Chinchipe, Ecuador; N. Krabbe). H: watkinsi (Pando, Bolivia; T. S. Schulenberg). I: possibly the unnamed taxon referred to by Remsen (2003) (Amazonas, Brazil; C. A. Marantz). J: rufipectus (Magdalena, Colombia; N. Krabbe). K: Hylocryptus erythrocephalus (Loja,Ecuador;N.Krabbe).L:Hylocryptusrectirostris (MinaisGerais,Brazil;H.G.Remold). 6 A B Z 1 1 3 lDU O iik ma 2 III ^ 1 1 1 4 TIME(sec.) Figure 3. Songs of three foliage-gleaners. A: Automolus rubiginosus brunnescens. B: A. r. rufipectus. C: Hylocryptuserythrocephalus. SameasFig.2G,JandK,butwithadifferenttimescale. 6 A A B B c c D D E E ^ 5 ^ 4 If II 1 1 li Hi i! 1 - 1 I \ ! t \ \ li 1 1 2.5 TIME(sec.) Figure 4. Calls of four taxa of foliage-gleaners. A: Automolus rubiginosus brunnescens (Napo, Ecuador; N, Krabbe).B:A. r. nigricauda(Esmeraldas,Ecuador;O.Jahn).C:Hylocryptuserythrocephalus(Piura,Peru;H.van Oosten).D-E:Automolusrubiginosusrufipectus(Magdalena,Colombia;N.Krabbe),Dforagingcalls,Ealarm- calls.Notetimescale. ridge,apparentlyoccurringwherevertherewasavailablehabitat.Thetwoclosestterritories found werec.200 mapart. Todd & Carriker (1922) noted Itwasperhaps mostnumerousat PuebloViejo in the thickscrub,butcannotbeconsidered a commonbird anywhere'. Large , NielsKrabbe 223 Bull. B.O.C. 2008 128(4) areas around Pueblo Viejo have been denuded by burning, but considerable expanses of forest and scrub occur between Pueblo Viejo and the San Lorenzo ridge. In view of the species' tolerance of disturbed habitats, there seems to be no grounds for considering it immediately threatened. It is probably best given the lUCN status of Near Threatened, approachingthe status ofVulnerable (criteria Bl(b), B2(b) and CI). Tomyknowledge, only a smallpartofits rangeis effectivelyprotected (the 'ElDorado' reserve onthe SanLorenzo ridge). The Sierra Nevada de Santa Marta National Park only includes higher elevations in this part of the mountains, and nearby Tayrona National Park only elevations below the range ofrufipectus. Discussion A detailed study and analysis mayreveal the small differencesbetween songs ofsome ofthe othersubspecies ofAutomolus rubiginosiis tobe constantand perhaps sufficientlydif- ferent for species recognition (see Remsen 2003), were they ever to come into contact. However, theirclose similarityto each other suggests that they likely form a monophyletic unit. In contrast, lesser vocal differences between A. rufipectus and H. erythrocephalus sug- gestthatA. rufipectusmaynotbeacloserelativeofitsproposedcongenersandconspecifics. The habitat of rufipectus is distinctly drier than those of other forms referred to A. rubigi- nosus. However, in absence of a study of molecular or other characters, convergent evolution cannotbe ruled out. Independent of whether rufipectus is a close or distant relative of other taxa currently placed in A. rubiginosus, the dramatic vocal differences demonstrate that rufipectus merits species recognition. The two sympatric species A. infuscatus and rufipileatus differ in fewer vocalvariables (Fig. 5a-c), indicatingthatevensmallvocaldifferencesmaybesufficientfor speciesrecognitionin foliage-gleaners. I therefore recommend thatspecies rankbe restitut- ed for rufipectus. Santa Marta Foliage-gleaner (rather than Rufous-chested Automolus as employedby Cory & Hellmayr 1925) mightbe an appropriate vernacularname. Vocalisations of the other six species referred to Automolus (Figs. 5a-g, 6a-c, 7a) are highly diverse and do not, in general, offer clues as to relationships among species. Only songs of infuscatus and rufipileatus (Fig. 5a-c) bear sufficient resemblance to each other that theycould be confused by experienced field ornithologists. Inter-relationships among the ten presently recognised genera of foliage-gleaners and allies are unclear, and several genera are so poorly defined that the the only solution to avoid problems in defining them morphologically would be to merge them all (Remsen 2003). In relation to other furnariids, the multiple-gene molecular studies by Irestedt et al. (2006; unpubl. data) provide strong evidence that foliage-gleaners and allies (including Megaxenops and Heliobletus) form a monophyletic unit; within the foliage-gleaner assembly these authors found two clades, one comprising Automolus/Hyloctistes Hylocryptus/Clibanornis and Thripadectes, and the other consisting of Philydor, Syndactyla, Anabacerthia, Heliobletus andMegaxenops. Genus Hylocryptus was erected for erythrocephalus when that species was described by Chapman(1919),mainlyonthebasisofitslong,straightbill. Hellmayr(inCory&Hellmayr 1925) left rectirostris inAutomolus, butnoted that itsjuvenile plumage and certain structur- al characters suggest affinities with erythrocephalus. Despite the widely disjunct distributions of the two species, Zimmer (1936) had 'no hesitation' in assigning rectirostris toHylocryptus,partlybecauseofitssimilargeneralcolourpatterntoerythrocephalus,butpri- marily because its bill, though shorter, approaches that of erythrocephalus in length and shape. Zimmer's treatment has been followed by all subsequent authors. Songs of the two NielsKrabbe 224 Bull. B.O.C.2008 128(4) A B C D E G _ 5 N I 4 ILU 3 iiiii 1 i tiKk't- 8m 2 . #.is f ' ..i„-r, a. li- 1 10 13 TIME(sec.) Figure5.SongsofdifferentAutomolusfoliage-gleaners.A:A. rufipUeatusconsobrinus (Sucumbios,Ecuador;J. V. Moore). B:A. infuscatus badius (Amazonas,Venezuela;C. Parrish).C:A. i. turdinus (Orellana,Ecuador;R. S. Ridgely).D:A.paraensis (MatoGrosso,Brazil;C.A.Marantz). E:A. leucophthalmuslammi(Alagoas,Brazil; C. A. Marantz). F: A. I. sulphurascens (Rio de Janeiro, Brazil; L. P. Gonzaga). G: A. melanopezus (Orellana, Ecuador;R.S.Ridgely). 6 A B c D E N I ^ 4 ft Z 3- fit)] UDJ O .filHIItffll LU CC LL 1 10 TIME(sec.) Figure 6. Songs of different Automolus and Anabazenops foliage-gleaners. A: Automolus ochrolaemus cervinigularis (Veracruz, Mexico; L. Irby Davis). B: Automolus o. exsertus (Costa Rica; L. Irby Davis). C: Automolus0. turdinus(Sucumbfos,Ecuador;P.Coopmans).D:Anabazenopsfuscus(RiodeJaneiro,Brazil;T.A. Parker). E:Anabazenopsdorsalis (MatoGrosso,Brazil;M. D. Medler). 10^ 8- 'n i 6- CY Z LDU 4 O UJ i f FR 2 H MnUfi iI t t i I 0- 10 13 TIME (sec.) Figure7. SongsofdifferentAutomolus and Syndactylafoliage-gleaners. A:A. r. roraimae (Bolivar,Venezuela; M.VanBeirs)(speciesremovedtoSyndactylabyZimmeretal.2008).B:S.ruficollis(Loja,Ecuador;N.Krabbe). C: S. subalarismentalis (Napo,Ecuador;M.Lysinger).D: S.g.guttulatus (Aragua,Venezuela;C.A.Marantz). E: S. rufosuperciliata cabanisi (Zamora-Chinchipe, Ecuador; N. Krabbe). For sonograms of S. dimidiata see Robbins&Zimmer (2005).Notefrequencyscale. NielsKrabbe 225 Bull. B.O.C.2008 128(4) species have rather different patterns, but show some similarity in pitch and quality (Fig. 2k-l). The resemblance of the song of rufipectiis to that of the much longer and straighter billed Hylocryptiis erythrocephalns (Figs. 2j-k, 3b-c) might be coincidental, as perhaps also suggested by their different calls (Fig. 4c-e), but is so striking, that the possibility that they aresistertaxa demands consideration. Were such a relationship confirmed,itwould add to the already interesting evolutionary and biogeographic history of the genus Hylocryptiis (whichwouldcomprisethreerelictualspecieswithdisjunctdistributionsinwidelyseparat- ed dry-forest areas of the Neotropics). It would also show that songs of foliage-gleaners, even in relatively smallpopulations, may remain unchanged for extended periods oftime, andwouldcallforadditionalcautionwhenusingbillshapeasataxonomiccharacterinthis group. A similar case ofvocal resemblance in songbetweenforms whose morphologies do not suggest that they are closely related has been reported between the fairly straight- mandibled (and rather uniform) Automolus roraimae (Fig. 7a) and the distinctly recurve-mandibled (andmoreorlessstreaked) members ofthegenus Syndactyla (Fig. 7b-e) (seealsoHilty2003, Robbins & Zimmer2005,Zimmeretal. 2008). Inview ofthehighlydis- puted merging ofmanyfoliage-gleaner generabyVaurie (1980) (see Remsen2003, Robbins & Zimmer 2005) and the conflicting indications of morphological and vocal data as to the relations ofthe taxon rufipectus, I would propose awaiting the results of a molecular study before making a generic change for this foliage-gleaner. The Sierra Nevada de Santa Marta is an isolated massifin northern Colombia with an area ofc.5,900 km^. Withno fewerthan70 endemicbird taxa, itisperhapsthe mostdiverse smaller centre of terrestrial endemism in the world. Most of the endemic taxa are derived from Andean forms (Todd & Carriker 1922), and their many different levels ofdifferentia- tion suggest that multiple colonisations have occurred. Seventeen are presently ranked as speciesbyRemsenetal (2007). Manyofthesespeciesareingroupswithcolourfulplumage, whilst many suboscines remain ranked only as subspecies. It therefore comes as no great surprise that A. rufipectus is better ranked as a biological species. Preliminary analyses of vocalisations suggest that species rank is likely to be appropriate for several other taxa endemic to the Santa Marta mountains, such as Trogon personatus sanctaemartae, Lepidocolaptes lacrymiger sanctaemartae, Grallaria rufula spatiator, Henicorhina leucophrys ana- choreta,Arremon torquatusbasilicus (alreadyshowntobegeneticallydistinctbyCadena2006, Cadena etal. 2007), and possibly others. Acknowledgements I thankC. D. Cadena, M. B. Robbins,J. Fjeldsa,J. V. Remsen, A. M. Cuervo and T. M. Doneganfor useful commentsonthemanuscript.ThefieldworkwaspartiallysponsoredbyProAves,Colombia. References: Bangs,O. 1898.OnsomebirdsfromPuebloViejo,Colombia.Proc. Biol. Soc. Wash. 12: 157-160. Boesman,P. 1999.BirdsofVenezuela.Photographs,soundsanddistributions.CD-ROM.BirdSongsInternational, Westernieland. Boesman,P. 2007. BirdsofVenezuela. DVD-ROM. Birdsounds.nl,Winsum. Cadena,C.D.2006.BiogeographyofBuarremonbrush-finches(Aves,Emberizinae):integratingecology,evo- lution,andsystematics.Ph.D. dissertation.Univ.ofMissouri,StLouis. Cadena, C. D., Klicka, J. & Ricklefs, R. E. 2007. Evolutionary differentiation in the Neotropical montane region: molecular phylogenetics and phylography of Buarremon brush-finches (Aves: Emberizidae). Mol. Phyl. &Evol.44:993-1016. Chapman, F. M. 1919. Descriptions ofproposed newbirds from Peru, Bolivia, Brazil, and Colombia. Proc. Biol. Soc. Wash. 32:253-268. Coopmans,P.,Moore,J.V.,Krabbe,N.,Jahn,O.,Berg,K.S.,Lysinger,M.,Navarrete,L.&Ridgely,R.S.2004. 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ThebirdsoftheSantaMartaregionofColombia: astudyinaltitudinal distribution.Ann. CarnegieMus. 14. Vaurie, C. 1980. Taxonomy and geographical distribution of the Furnariidae (Aves, Passeriformes). Bull. Amer.Mus. Nat. Hist. 166: 1-357. Vaurie, C. & Schwartz, P. 1972. Morphology and vocalizations of Synallaxis unirufa and Synallaxis castanea (Furnariidae,Aves),withcommentsonotherSynallaxis.Amer.Mus. Novit. 2483. Whitney, B. M. & Pacheco, J. F. 1994. Behavior and vocalizations of Gyalophylax and Megaxenops (Furnariidae),twolittle-knowngeneraendemictonortheasternBrazil. Condor96:559-565. Zimmer, K. J. 1997. Species limits in Cranioleuca vulpina. Pp. 849-864 in Remsen, J. V. (ed.) Studies in NeotropicalornithologyhonoringTedParker.Orn.Monogr.48. NielsKrabbe 227 Bull. B.O.C.2008 128(4) Zimmer,K.J.2002. Specieslimitsinolive-backedfoliage-gleaners{Aiitomolus;Furnariidae). Wilson Bull. 114: 20-37. ZinrmerK.J.,Robbins,M.B.&Kopuchian,C.2008.Taxonomy,vocalizations,syringealmorphologyandnat- uralhistoryofAntomolusroraimae(Furnariidae). Bull. Brit. Orn. CI. 128: 187-206. Address: Zoological Museum, University ofCopenhagen, Universitetsparken 15, DK-2100, Copenhagen 0, Denmark,e-mail:[email protected] ©BritishOrnithologists'Club2008 APPENDIX Songsexamined.Abbreviationsusedare:XC=Xeno-cantowebsiteatwww.xeno-canto.org;LNS=Macaulay LibraryofNaturalSounds,Cornell,websiteatwww.animalbehaviorarchive.org/loginPublic.do.Recordings knowntobegivenbyanindividualalreadyrepresented,wereavoidedorcountedasone. Automolus rubiginosus: A. r. rubiginosus 6 (Boesman2007; LNS89684, 95077, 127210, 127230, 127246, 127247, 127271);A. r. veraepacis 3 (Boesman2007;XC 11012, 14140);A. r. saturatus 1 (LNS 108866);A. r. nigricauda 9 (Jahnetal. 2002, Krabbe «&Nilsson2003; XC 5576, 6598, 8000, 8308 (call), 13139);A. r. sasaimae 1 (XC 12728, 12729,12731);A. r.obscurus1 (XC7520);A. r. brunnescens4(Krabbe&Nilsson2003;XC6575;alsoanunpubl. recording of calls by N. Krabbe); A. r. watkinsi 1 (Schulenberg et al. 2000); A. r. ssp. nov.: 1 (LNS 117196, 117197). Automolus rufipectus: 6 (Boesman 1999; Strewe et al. 2003; also songs of four individuals and calls of one recordedduringthepresentstudy). Automolusroraimae:3(Boesman2007). Automolusochrolaemus:A.o. cervinigularis5(LNS7009,7011,7012, 102599;XC 14133);A.o.hypophaeus2(LNS 7012;XC11011);A.o.exsertus1 (LNS7013);A.o.pallidigularis11 Qahnetal.2002,Krabbe&Nilsson2003;LNS 7014, 82449; XC 1490, 12898); A. o. turdinus 12 (Krabbe & Nilsson 2003, Moore 1993; XC 6094, 8704; also 7 unpubl. recordingsfromEcuadorbyP. CoopmansandJ. V. Moore);A. o. ochrolaemus5 (LNS39003;XC 85, 8361,8538, 13424);A. o. auricularis4(LNS88948;XC9540,9541, 14773). Automolus infuscatus:A. i. infuscatus 15 (Boesman2007,Krabbe&Nilsson2003,Moore1993;XC3434, 10324; LNS53363,53377,110437;also4unpubl.recordingsfromEcuadorbyP.Coopmans,M.Lysinger,J.V.Moore andR.S. Ridgely);A. i. badius4(Boesman2007;LNS117037;XC6229). Automolus paraensis (see Zimmer 2002): 11 (LNS 88849, 88900, 88950, 89066, 89067, 106198, 110005, 113065, 115019, 115233, 115237). Automolus leucophthalmus: A. I. lammi 3 (LNS 127914, 127950, 127992); A. I. sulphurascens 10 (Gonzaga & Castiglioni2001;LNS19082, 19086, 19214, 113312, 113394, 113395, 115354, 115426, 115429). Automolusrufipileatus:A. r. consobrinus22(Krabbe&Nilsson2003,Moore1996;LNS115874, 119700, 110635, 106422, 110523, 129499, 129534; XC 222,3206,6409,9926, 13425;also4unpubl. recordingsfromEcuadorby J. V. Moore and R. S. Ridgely); A. r. rufipileatus 8 (LNS 88386, 88462, 88522, 88634, 88668, 106104, 10630, 109948). Automolusmelanopezus: 13 (Krabbe&Nilsson2003;LNS26565,38862,38865,38885,38914;XC 12027;Moore 1993;also5unpubl.recordingsfromEcuadorbyP.Coopmans,M. Lysinger,J.V. MooreandR.S. Ridgely). Hylocryptus erythrocephalus: 11 (Coopmans et al. 2004, Krabbe & Nilsson 2003; LNS 122729; XC 4152, 4191, 8306,8308(call),8727, 14414). Hylocryptusrectirostris: 2(LNS114856;XC 11924). Anabazenopsfuscus: 8(Gonzaga&Castiglioni2001;LNS39082, 103868, 112723;XC4861,4862,5468, 14680). Anabazenopsdorsalis: 8(LNS88418,106182, 106092, 106175, 106200, 109946;XC6574, 11902). Syndactylaguttulata: S.g.guttulata6(Boesman2007;LNS127816, 127819). Syndactylaruficollis:6(Coopmansetal.2004,Krabbe&Nilsson2003). Syndactylarufosuperciliata:S. r. rufosuperciliata2(Gonzaga&Castiglioni2001;LNS108238);S. r.acrita2(LNS 19094; XC 8195); S. r. oleaginea 1 (Krabbeetal. 2001); S. r. cabanisi3 (Lysingeretal. 2005, Krabbe & Nilsson 2003). Syndactylasubalaris: S. s. mentalis3(Moore&Lysinger1997,Krabbe&Nilsson2003);S. s.subalaris3(Krabbe &Nilsson2003).