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The Auk 116(4):891-911, 1999

PHYLOGENETIC ANALYSIS OF THE NEST ARCHITECTURE OF NEOTROPICAL (FURNARIIDAE)

KRZYSZTOF ZYSKOWSKI • AND RICHARD O. PRUM NaturalHistory Museum and Department of Ecologyand Evolutionary Biology, University of Kansas,Lawrence, Kansas66045, USA

ABSTRACT.--Wereviewed the tremendousarchitectural diversity of (Furnari- idae) nestsbased on literature,museum collections, and new field observations.With few exceptions,furnariids exhibited low intraspecificvariation for the nestcharacters hypothe- sized,with the majorityof variationbeing hierarchicallydistributed among taxa. We hy- pothesizednest homologies for 168species in 41 genera(ca. 70% of all speciesand genera) and codedthem as 24 derivedcharacters. Forty-eight most-parsimonious trees (41 steps,CI = 0.98, RC = 0.97) resultedfrom a parsimonyanalysis of the equallyweighted characters using PAUP,with the Dendrocolaptidaeand Formicarioideaas successiveoutgroups. The strict-consensustopology based on thesetrees contained 15 cladesrepresenting both tra- ditionaltaxa and novelphylogenetic groupings. Comparisons with the outgroupsdemon- stratethat cavitynesting is plesiomorphicto the furnariids.In the two lineageswhere the primitivecavity nest has been lost, novel nest structures have evolved to enclosethe nest contents:the clayoven of Furnariusand the domedvegetative nest of the synallaxineclade. Althoughour phylogenetichypothesis should be consideredas a heuristicprediction to be testedsubsequently by additionalcharacter evidence, this first cladisticanalysis of the fur- nariids demonstratesthe generalutility of nest charactersin reconstructionof avian rela- tionships,and it providesa testof monophylyfor several furnariid taxa. Received 29July 1998, accepted12 March 1999.

RECENT APPLICATIONS OF PHYLOGENETIC SYS- a broad diversityand complexityof neststruc- TEMATICSto ethologyhave demonstratedthat tures. behavioraltraits can be as historicallyinfor- The suboscinefamily Furnariidae,with 240 mativeas morphologicaland molecularchar- currently recognized biological , is acters(Prum 1990, de Queiroz and Wimberger amongthe mostmorphologically, behaviorally, 1993,Paterson et al. 1995,Irwin 1996,Kennedy and ecologicallydiverse families of et al. 1996).Nests are a detailedpart of the ex- (Ridgelyand Tudor 1994, Skutch1996). The tendedphenotype of ,and they providea furnariid radiationincludes phenotypes con- durablephysical record of the behaviorthat re- vergenton numerousfamilies of oscines,such sultedin their construction(Hansell 1984, Col- aslarks, jays, tits, creepers,nuthatches, wrens, lias 1986).A wealth of quantifiabledetail in thrushes, thrashers, dippers, and warblers neststructure, placement, and ontogenymakes (Leisler1977). Representatives of the Furnari- nests amenableto comparativeanalysis. Nest idae can be foundin all Neotropicalhabitats, architecturecan provide evidence of phyloge- fromcoastal surf zone and sand dunes through netic relationships(Kiff 1977, Lanyon 1986, dry andhumid lowland and montane to Prum 1993,Whitney et al. 1996).Furthermore, the treelesspuna of the . hypothesesof phylogenycan be used to dis- The diversityof nestplacement and structure coverhistorical patterns in the evolutionof nest in the Furnariidaeapproaches that of theentire featuresand to test hypothesesabout adapta- order Passeriformes(Narosky et al. 1983,Sick tion in nest design (Dekker and Brom 1992, 1993,Collias 1997).Nests can be placedin ex- Winklet and Sheldon 1993, Lee et al. 1996, cavatedor adoptedcavities under or abovethe Eberhard1998, Sturmbauer et al. 1998).Here, ground. Non-cavity nests can be supported we use nest architectureto generatea hypoth- from below,attached by the side to a vertical esisof phylogeneticrelationships for the Neo- support,or suspendedfrom a branchor rock. tropicalovenbirds (Furnariidae), which exhibit Nest structuraldiversity ranges from shallow pads of looselypiled material to domednests E-mail: [email protected] with constructed roofs of interlaced sticks, in-

891 892 Z¾SKOWSK•AND PRUM [Auk, Vol. 116 terwoven soft vegetation, or clay. Additional nariidae + Dendrocolaptidaeis a clade com- elaborations of domed nests include entrance posedof the ,Conopophagidae, tubes, tunnels with constrictions,awnings and Rhinocryptidae(i.e. Formicarioideasensu above the entrance,nonbreeding chambers, Sibleyand Ahlquist1990). and adornmentwith conspicuousobjects. The genericlimits within Furnariidaerec- Over the last century,architectural features ognizedin this study follow thoseof Peters of furnariidnests have been recognized to be (1951), with a few modificationsfollowing evolutionarilyconservative, constituting poten- Ridgely and Tudor (1994) and Kratter and tially informativesystematic characters (Iher- Parker(1997). Since the publicationof Peters' ing 1914, 1915; Vaurie 1971, 1980). However, (1951)volume, a surprising14 new speciesand previousauthors have used nest architecture in 1 new genusof furnariidshave been described, an eclectic,noncladistic manner; no compre- 14 subspecieshave been elevated to therank of hensivephylogenetic analysis of thefurnariids species,and 5 specieshave been synonymized hasbeen conducted, preventing rigorous study (Ridgelyand Tudor 1994,Pacheco and Gonza- of the evolution of nest-construction behavior. ga 1995,Silva 1995,Pacheco et al. 1996,Maijer Detailedhypotheses of hornologyamong nest and Fjelds• 1997, Zimmer 1997).As currently typeshave proven difficult to developbecause recognized,the Furnariidae consists of 240spe- of limited information about many species, ciesin 59 genera. paucityof behavioralinformation on the steps of nestbuilding (i.e. ontogeny),and the histor- METHODS ical lackof a phylogeneticmethodology. Data.--Variation in nest architecture of furnariids Our objectivewas to examinepatterns of var- iation in nest structure and construction behav- and outgrouptaxa was reviewedbased on the sci- entific literature and on data associated with nest ior within theFurnariidae and to performa cla- and eggspecimens in majorcollections. The first au- disticanalysis of the family basedon nestchar- thor examinednest specimens in majorNew World acters.We performedthis analysisto yield hy- collections(see Appendix 1) and collectednew data pothesesof phylogeneticrelationships of major describing204 nestsfrom 53 speciesduring his field cladeswithin the family and to evaluatethe studiesin ,, Paraguay,and (1994 usefulnessof nestcharacters for reconstructing to 1998).Results of the field work, includingnew on- avian phylogenies.We also proposeand dis- togeneticinformation and nest descriptionsfor sev- cussnew hypothesesregarding the evolution en specieswith previouslyundescribed nests, will be of furnariid nest-construction behavior publishedseparately. Nest informationwas gatheredfor 184 furnariid Reviewof furnariid systematics.--The Furnari- speciesfrom 50 genera;species for which nest de- idaeis a family of tracheophonesuboscine pas- scriptionswere demonstrablyerroneous or lacking serinesclosely related to the criticaldetails were subsequentlyexcluded. Data for (Dendrocolaptidae).The two familiesshare a the remaining168 species,representing 41 genera unique intrinsic syringeal muscle, M. vocalis (ca.70% of all speciesand genera),were used in the dorsalis(Ames 1971). Monophyly of the Den- analysis(Appendix 2). A completelist of sourcesof drocolaptidaehas been supported by morpho- nestdescriptions for eachspecies is presentedin Ap- logicalsynapomorphies (Raikow 1994, Clench pendix1. Samplesizes varied considerably, with ap- proximately5% of the speciesrepresented by more 1995), but no morphologicalsynapomorphy than 50 nestseach, the majorityof speciesbased on satisfactorilydemonstrates the monophylyof more than 10 nests,and about 20% of the species the furnariids.Morphological characters tradi- known from a singlenest description. tionally used to diagnosethe family (e.g.lack We also assemblednest informationfor outgroup of hornson the processivocales [Ames 1971] taxa: 37 speciesrepresenting all 13 generaof Den- andpseudoschizorhinal nares [Feduccia 1973]) drocolaptidae;19 speciesrepresenting 8 of 12 genera are primitive amongtracheophones, and none of Rhinocryptidae;19 speciesrepresenting all 7 gen- is shared by all furnariid species(Feduccia era of Formicariidae;and 5 of the 8 speciesof Cono- pophagidae. 1973).We assumedthat the Furnariidaeand the Phylogeneticanalysis.--The number of ingrouptaxa Dendrocolaptidaeare eachmonophyletic and was reduced to 49 operational taxonomic units are sistertaxa, basedon the Sibley and Ahl- (OTU) by combiningcongeneric species having iden- quist(1990) DNA-DNA hybridizationhypoth- tical characterstates. Appendix 2 includesdefini- esis.The hypothesized sister group to theFur- tions of speciesgroups composedof behaviorally October1999] NestPhylogeny ofFurnariidae 893 similarcongeners used as OTUs. The outgroupspe- data that are consistent with those of their bet- cies were condensedinto two OTUs representing ter knowncongeners are presumed to be mem- two successiveoutgroups, Dendrocolaptidae and bersof the sameOTU (seeAppendix 2). Formicarioidea. Webased hypotheses of behavioralhomologies on 1. Nest in cavity.--Thenest is placed in an specialdetailed similarities in neststructure and on- adoptedor self-excavatedcavity, such as a sub- togenyof nestconstruction. Variations in nestarchi- terranean burrow, rock crevice, termite mound, tecture were coded as 24 characters, of which 22 were tree cavity,bamboo internode, cavity in a man- binary and two trinary. Apparentautapomorphies made structure, or enclosed construction of were included in the data set because these charac- ters are synapomorphiesof polytypicterminal taxa otherbirds or mammals.Nests placed in cavi- (characters4, 10, 20, 24). Genuineautapomorphies ties are presentin ,, Au- were alsoincluded because they may be sharedby tomolus, Chilia, ,Furnarius, , unidentifiedcryptic species within current species or ,Hyloctistes, Lochmias, Ochetorhyn- shared with behaviorallyunknown taxa and ulti- chus,, Pseudocolaptes, Pygarrhichas, Scle- matelybe phylogenetically informative (characters 3, rurus,, , and .In 5, 8, 12, 19,22). Characterpolarities were inferred by addition,we codedas polymorphic(0 and 1) outgroupcomparisons. In instanceswhere the sec- the Asthenespyrrholeuca group, Leptasthenura, ond outgroupwas polymorphic, we usedpresumed andPremnoplex brunnescens, because these spe- phylogeneticrelationships within the Formicarioi- ciesplace some nests in cavitiesand others in deato inferthe primitivecharacter state for the out- groups.Taxa exhibiting both ancestral and derived the open. Among the outgroup taxa, cavity statesof a givencharacter were coded as polymor- nestingis presentin all known dendrocolap- phic(0 and 1) for that character.All characterswere tids and in several formicarioids. weightedequally. 2. Formof cavitynest.--We hypothesize two The most-parsimoniousphylogenetic resolutions ordered derived states of nests built in cavities. of the nest data were identified using PAUP 3.1.1 (2.1)Non-pliable plant material is eitherpiled (Swofford1993). We used a constrainttree assuming or looselyinterlaced to form a loose,usually monophylyof the ingroupand the outgroupstruc- unlinedplatform (Fig. 1A). Platformsare pres- ture describedabove. We performed 100 replicate heuristic searches with random addition of taxa to ent in the Automolusleucophthalmus group, Chi- lia, Hyloctistes,Pseudocolaptes, Pygarrhichas, minimizeinput order bias and ensure unbiased sam- pling of tree space.The tree bisection-reconnectionSclerurus, and Thripadectesmelanorhynchus. (2.2) algorithmwas employed, and zero-lengthbranches Pliableplant material is wovenand compacted werecollapsed to yield polytomies.We used a strict- into a shallowcup, often lined with nonvege- consensustree to summarizethe set of maximally tative materialsuch as wool or feathers(Figs. parsimonioustopologies identified by PAUP.Char- 1B-D). Cups are presentin Anabazenops,Auto- acter evolutionwas analyzed by examiningparsi- molusrubiginosus, Cinclodes, Furnarius, Geositta, moniouscharacter optimizations on fundamental Hylocryptus, certhioides, Philydor, treesusing MacClade 3.03 (Maddison and Maddison Thripadectesrufobrunneus group, Upucerthia, 1992). and Xenops.Cup-like nestsof Aphrastura,Lep- tasthenura,Spartonoica, and someAsthenes are CHARACTER ANALYSIS hypothesizedto be nonhomologousbecause they representreduced domed constructions Each nest character below includes a brief de- (seecharacter 11). scriptionof the derivedstate and its taxonomic Althoughboth platforms and cupsare found distribution.Aspects of nest architectureana- in the two successiveoutgroups, the cuphas a lyzed includenest form and structure,materi- narrower taxonomic distribution and therefore als, nest location and attachmentto substrate, is hypothesizedto be independentlyderived. and details of construction behavior The first For example,when cup and platformare su- nine charactersdescribe nests placed in cavities perimposedon the dendrocolaptidphylogeny (Fig. 1), whereasthe remaining15 characters of Raikow(1994), platforms are shared by most focuson non-cavitynests (Fig. 2). The distri- lineagesincluding the most basal whereas butionof characterstates in bothingroup and cups appear to have been derived indepen- outgrouptaxa is given in Appendix2. Each dentlyin two lineages.Platforms of barkflakes OTU is basedupon at least one behaviorally or dry leavesshared by mostdendrocolaptids, well-knownspecies. Species with partial nest but not foundin the ingroup,are assumedto 894 ZYSKOWSKIAND PRUM [Auk, Vol. 116

B

FIG. 1. Structureof furnariid nestsplaced in cavities.(^) Hyloctistessubulatus: platform of compound- leaf rachises(photo by IN.Wheelwright). (B) Cinciodesexcelsior: cup of dry grass(WFVZ 162881).(C) Thri- padectesvirgaticeps: cup of rootlets(WFVZ 159222).(D) Furnariusrufus: cross-section of clay oven revealing grasscup (KU 89623).(E) Furnariuscristatus: day oven(KU 89619).(F) Lochrniasnematura: domed nest of bamboo leaves (KU 89612). Nests ^, B, C, and F were extractedfrom burrows. be ancestralto the furnariid platformscoded membraneous tree-fern scales (ramenta) is below as characters 3 to 6. piled inside a cavity and is sometimesused to 3. Platformof woodchips.--Eggs rest directly cover the eggs on departure of incubating on wood chipsfrom cavity excavation,occa- birds. Ramentaare used exclusivelyby Pseu- sionallysupplemented with sparseplant ma- docolaptes. terial or down. We hypothesizethat this ex- 5. Platformof sticksand feathers.--A fairly tremelyreduced cavity lining is derivedfrom a massiveplatform is constructedof dry sticks dendrocolaptid-like platform. Platforms of and feathersof larger birds (usually rectrices wood chipsgenerated passively during cavity and remigesthat havea distinctlystiff rachis). excavationare characteristicof Pygarrhichas. This characteris presentin Chilia. 4. Platformof fern scales.--Athick layer of 6. Platformof leaf rachises.--A platform is con- October1999] NestPhylogeny ofFurnariidae 895

D - E F

FIC. 2. Structureof non-cavitynests of furnariids.(A) Phleocryptesmelanops: domed nest with awning abovethe entrance,attached to verticalstems. (B) Leptasthenuraandicola: domed nest of dry grass(USNM 30597).(C) Premnoplexbrunnescens: domed nest of moss,suspended from a log (photoby M. Marin). (D) Phacellodomusdorsalis: secondarily pensile nest of sticks(photo by J.O'Neill). (E) Asthenesbaeri: nest of sticks saddledon a cactus.(F) Schoeniophylaxphryganophila: restricted lining of conspicuousmaterial, inside a domedstick nest (KU 89629).Arrows indicatenest entrances where necessary.

structedentirely of compound-leafrachises, built of wiry plant materialsuch as rootlets, usuallyof a singleplant species(Fig. 1A). This root fibers,or fungal rhizomorphsof Maras- characteris presentin the Automolusleuco- mius(Fig. 1C) is presentin Automolusrubigino- phthalmusgroup, Hyloctistes,, and sus,Hylocryptus, Philydor, Thripadectes rufobrun- Thripadectesmelanorhynchus. neusgroup, and Xenops. 7. Cupcomposition.--We hypothesize two un- 8. Profuselining of snakeexuviae.--A large ordered derived states for cup composition. quantity of snakeexuvia or human-madema- (7.1) A cup constructedof ribbon-likevegeta- terial of similar appearance(e.g. cellophane) tive material suchas grassleaves and stripsof linesa vegetativecup. This characteris present inner bark (Figs.lB,D), is presentin Anabazen- in Ochetorhynchuscerthioides. ops, Cinclodes,Furnarius, Geositta, Ochetorhyn- 9. Tree-cavityexcavation.--The birds them- chuscerthioides, and Upucerthia.(7.2) A cup selvesexcavate a cavityin softwood. Pygarrhi- 896 Z¾SKOWSK•^NI• PRVM [Auk, Vol. 116

chasand Xenopsare the only furnariidsknown ensupon drying, are woven into the nestwall. to excavatesuch cavities, but even they occa- Mud-daubingis known only in Phleocryptes. sionallyadopt existing holes. 13. Outer shellof stems.--A vertically ovoid 10. Clay dome.--Anigloo-shaped structure, nest of fresh Sphagnummoss is looselyencap- often described as an "oven," is constructed of sulatedby herbaceousstems. This characteris claymixed with plant fibers,hair, or dung(Fig. presentin Schizoeacaand Oreophylax. 1E). The wall at one side of the entranceis fold- 14. Sticks.--Bulkof domednest is composed ed inwardto dividethe globeinto two helically of dry and oftenthorny, woody sticksthat are arrangedchambers: an inner incubationcham- tightly interlaced(Figs. 2D-F). In human-al- ber, and an outer antechamber;only the inner teredhabitats, sticks can be replacedwith wire, chamberis lined (Fig. 1D). Clay-ovenconstruc- including barbed wire (Sick 1993:434).Present tion has beenreported from all speciesof Fur- in Acrobatornis, Anumbius, Asthenes humicola narius,but somepopulations of F.figulus may be group,Asthenes anthoides, some Asthenes pyrrho- obligateadopters of existingovens and other leucagroup (codedas polymorphic), , cavities (Studer and Vielliard 1990, A. Whitta- Coryphistera,Eremobius, Gyalophylax, Phacello- ker pers.comm., K. Zyskowskipers. obs.). domus,Poecilurus, Pseudoseisura, Schoeniophylax, 11. Domed vegetativestructure.--Vegetative and .Although the Cranioleucapyr- material is used to construct a domed nest rhophiagroup is known to incorporatesticks with an entrance reduced to a hole on the side, duringnest construction, the bulk of thenest is top, or bottom of the nest (Figs. 1F and 2A-E). made of interwovenpliable materials, so we Domed vegetative nests are present in Acro- codedthis OTU as lackingthe derivedstate. batornis,Anumbius, Asthenes, Certhiaxis, Cory- 15. Extent of lining.--We hypothesizetwo phistera,, Eremobius, Gyalophylax, unordered derived states for the extent of stick Leptasthenura,, Limnornis, Lochmias, nestlining. (15.1)The entire or large portionof ,Oreophylax, , Phleo- a nestinterior is lined with soft plant material, cryptes, Poecilurus, brunnescens, feathers,or hair This extensivelining is pres- Pseudoseisura,Schizoeaca, Schoeniophylax, Sip- ent in Acrobatornis, Anumbius, Asthenes anthoi- tornis,, Synallaxis, and Thri- des, Astheneshumicola and A. pyrrholeuca pophagafusciceps; apparently, they are inde- groups,Coryphistera, and Phacellodomus.(15.2) pendently derived in severalrhinocryptids in A thin circularpad of soft, oftenbrightly col- the secondoutgroup. ored material (usually green,pubescent dicot- Cup-shapednests of Aphrasturaspinicauda, a yledonousleaves, but sometimesorange or cavity-nester,are probablyhomologous with white arthropodcocoons and oothecas)is re- domednests because their lining is not restrict- stricted to the bottom of the incubation cham- ed to the bottomof the cavitybut alsopartially ber (Fig. 2F). The restrictedlining is presentin coversits walls (Johnson1967:175). In support Certhiaxis,Gyalophylax, Poecilurus, Pseudoseisu- of this hypothesis,some species of Leptasthen-ra, Schoeniophylax,and Synallaxis.Some nests of ura and Asthenesconstruct fully domednests in Certhiaxis and Pseudoseisura are found with open situationsbut only much reduced cups eggsresting directly on an unlined floor of a (muchlike thoseof Aphrastura)when placed in- broodchamber, perhaps indicating that the lin- sidecavities. All three generaare codedas de- ing is added during incubation(e.g. Certhiaxis rived for this character. cinnamomea;K. Zyskowskipers. obs.).Species Similarly,Spartonoica construct rather rudi- of Pseudoseisuraaresignificantly larger than the mentary ground nests sometimesinterpreted remainingtaxa listed above,and they line their as open cups.Because these nests usually are nests with coarser material such as flakes of placedin a hollow in the baseof a clump of bark, small sticks,or dung. rushesor grass,the surroundingdense vege- 16. Thatch.--A varying quantity of sticks tation functionsas the roof and walls (Narosky and plant detritus is placed over the brood 1973).We hypothesizethat suchconstructions chamberforming a pile. Thatch is presentin are homologouswith domedvegetative nests, Certhiaxis,Gyalophylax, Poecilurus, Schoeniophy- and hence code them as derived. lax, and Synallaxis. 12. Mud-coatedvegetation.--Strips of macer- 17. Roofadornment.--Conspicuous objects of atedplant material coatedwith mud thathard- or human origin (e.g. whitenedbones, October1999] NestPhylogeny ofFurnariidae 897

brightlycolored or shinypieces of plasticor Nestsof othermarsh-nesting furnariids, es- glass,etc.) are placedon the roof and around peciallythose of Cranioleucasulphurifera, some- the entrance of a domed nest after it is com- times appearto be attachedto verticalstems. pleted. Presentin Anumbiusand Coryphistera. Thesespecies are not codedas derived,how- We are uncertain whether adornment with ever, becausein such instances the materials of largefeathers reported for Pseudoseisuralopho- the nestwall arenot attachedfirmly to upright tes(Nores and Nores1994) is homologous(cod- stems,and the actualnest support comes from ed "?"). the vegetationbelow the nest. In somecases, 18. Entrance tube.--Entrance to domed nest Cranioleucasulphurifera even use old Phleocryp- has a tubular extension that is either horizontal tesconstructions as a nestfoundation (Narosky or curvesupward. Entrancetubes are present et al. 1983). in Acrobatornis,Asthenes humicola group, some 23. Pensile nest.--Voluminous nest, with a Asthenespyrrholeuca group (coded polymor- small brood chamber entered from below, is phic),Certhiaxis, Eremobius, Gyalophylax, Phacel- constructedfrom the top down (cf. character lodomus,Poecilurus, Pseudoseisura, Schoeniophy- 24), presumablyby draping long strands of lax, and Synallaxis.We follow Narosky et al. green mossor strips of other plant material (1983:8)in consideringtubes not homologous from a branchor other woody or rocky over- with tunnels,which represent extensions of en- hang.Present in the Cranioleucaalbiceps group, trance passagesthat are entirely imbedded Margarornis,Premnoplex brunnescens, Siptornis, within a globularnest structure. and Thripophagafusciceps. 19. Double entrance.--Twoascending en- Nests of Premnoplexbrunnescens and Marga- trancetubes on oppositesides of a nest con- rornisappear homologous with pensilemoss vergeinto a singlesuperior brood chamber. Al- constructionsof the C. albicepsgroup because thoughthe tubesare largelyimbedded within they are alsobuilt largelyof moss,have a dis- the walls of a nest,they are tightlywoven and proportionatelysmall brood chamberentered well differentiatedfrom the vegetativestrands from below,and usually are attachedto an un- looselydraped over them (K. Zyskowskipers. dersideof a log or rocky overhang(Fig. 2C). obs.).This characteris presentin ThripophagaSome nests of P.brunnescens (e.g. WFVZ 157319 fusciceps. and 157401; see also Vaurie 1980:231) are even 20. Entrance-tube constriction.--Floor eleva- suspendedfrom hangingvines in a manner tion inside the tubular entrance results in for- very similarto that of the C. albicepsgroup. mation of a small antechamber(false nest) in 24. Secondarilypensile nest.--A boot-shaped front of the main chamber; both chambers are nest of sticks, with a horizontal tubular en- lined. Constrictedentrance passages are pres- trance,is built fromthe bottom up. Nestbegins ent in Phacellodomus. as a platform resting on a distal part of a 21. Entranceawning.--Semicircular awning branch,and usually (but not always)becomes of wovenplant material projects over the lateral pensileby progressiveaccumulation of mass entranceof a domednest (Fig. 2A). Suchelab- (Fig. 2D). Hanging stick nestshave been re- oration is present in Phleocryptes,Limnoctites, ported in all Phacellodomus.Even P. striaticollis, andLimnornis. We hypothesize that the awning with typicallynon-pensile nests (Narosky et al. of looselyprojecting sticks reported in some 1983),tends to placeits constructionson distal nestsof Cranioleucapyrrhophia (Narosky et al. parts of branches,where they occasionallybe- 1983)is not homologous. comepensile (e.g. Pereira 1938:fig. 29). 22. Attachment to vertical stems.--A nest is at- tachedby the sidesto verticalstems of reedsor RESULTS rushes (Fig. 2A). Ribbon-like vegetative strandsoriginating from nestwalls are tightly The 24 nestcharacters support 48 most-par- woundaround the supportingstems. Adhesive simonioustrees, each with a lengthof 41 steps, propertiesof mud broughtalong with macer- consistencyindex of 0.98, and rescaledconsis- ated plant material assurefirm attachmentof tencyindex of 0.97.The strict-consensustopol- the nest to the smooth surfaces of movable ogy based on these trees contains15 clades reeds.This modeof nestattachment is present (Fig. 3; multigenericlineages labeled as clades only in Phleocryptes. A to J). The distributionof characterssupport- 898 z¾s•ows•^Nr) PRUI• [Auk,Vol. 116

Chi/ia me/anura Geositta(9) Upucerthia(7) Ochetorhynchuscerthioides Cinclodes(11 ) A Furnarius(6) Philydor(4) Anabazenops(2) Automolusrubiginosus Hylocryptus(2) Thripadectesrufobrunneus group (2) Xenops (3) Pygarrhichasalbogularis Pseudocolaptes(2) Hyloctistessubulatus Automolusleucophthalmus group (3) Thr/padectesmelanorhynchu$ Sclerurus(6) L ochmias nematura Aphrasturaspinicauda Leptasthenura(9) D Schizoeaca(4) ß Oreophylaxmoreirae Sylviorthorhynchusdesmursii Limnornis curvirostris Limnoctites rectirOstris E Phleocryptesmelanops Spartonoicamaluroides Cranioleucapyrrhophia group (7) Cranioleucaalbiceps group (11) Thripophagafusciceps Siptornisstriaticollis Premnoplexbrunnescens Margaromis(2) Asthene$wyatti group (7) Asthenespyrrholeuca group (2) Astheneshum/cola group (11) Asthenes anthoides Acrobatornis fonsecai , Anumbius annumbi ß Coryphisteraalaudina Phacellodomu$(8) Eremobiusphoenicurus Pseudoseisura(3) Certhiaxis(2) Schoeniophylaxphryganophila Gyalophylaxhellmayri Poecilurus(2) Synallaxis(19) DENDROCOLAPTIDAE FORMICARIOIDEA F•c.3. Strictconsensus of48 most-parsimonious trees(length = 41, CI = 0.98,RC = 0.97)based on 24 nest-architecturecharacters. Dendrocolaptidae andFormicarioidea wereused as the successive outgroups. NodesAto J are multigeneric lineages diagnosed bynest characters. Number ofspecies isindicated inpa- rentheses.Species groups are defined in Appendix2.

ingthese clades is shown on one of the funda- supportedbya singlederived character. Clades mentaltrees (Fig. 4). The basal polytomy of the A and B are diagnosableby the wovencup consensustree includes six diagnosablegroups (character2.2) and the platform of leaf rachises with unresolvedinterrelationships. The largest (character6), respectively.Chilia is distin- of them(clade C) is composedof thefurnariids guishedby the stick-and-featherplatform thatbuild domed nests of plantmaterial. The (character5).Pygarrhichas isunique among the remainingfive groups(clades A andB, plus cavity-nestingfurnariids in thatit layseggs on Chilia,Pygarrhichas, andPseudocolaptes) include the floor of a tree cavity that remainsessen- speciesthat nest in cavities,asdo most of the tiallyunlined except for thepassive accumu- outgroup taxa. lationof woodchips (character 3). Pseudocolap- Eachof the five cavity-nestinglineages is tesis supportedby thebehavior of pilingup October1999] NestPhylogeny ofFurnariidae 899

Phleocryptes Sylviorthorhynchusiii•• Spartonoica '" L.c.Lt Aw

H.s. ' Sclerurus•,, tm'H•' Cran•?bl•eC•s

Thripadectes •:f',,• Anumbius virgaticepsceps Ph A.Ln•7.2• 13

Geositta • Astheneshumicola

Fumarius Phacellodomus

• sy. Pseudoseisura Ochetorhynchus• , c.m. Pygarrhichas• • Schoeniophylax Pseudocolaptes Certhiaxis

Fie. 4. One of 48 fundamentaltrees showing distribution of synapomorphiesand autapomorphiesand diagramsof selectednest designs.Characters with multiple optimizations(1, 2.1, and 9) are not shown. Numberscorrespond to nestcharacters in text, and namesof OTUs are abbreviationsof thosein the strict- consensustree (Fig. 3).

fern scaleson the bottomof a tree cavity(char- molusand Thripadectesare representedby dif- acter 4). ferent membersin cladesA and B; Hyloctistes Thewoven-cup clade (A) is composedof fur- and all Sclerurus are included in clade B. nariidsthat constructnests of grass-likemate- The diverseclade C, diagnosableby domed rial (Ochetorhynchuscerthioides and all behav- nestsconstructed of plant material includes11 iorally known speciesof Geositta,Upucerthia, basallineages with unresolvedrelationships. Cinclodes,Furnarius, and Anabazenops)and Three of theselineages, Lochmias, Aphrastura, thosethat build nestsof rootlets(Philydor, Hy- and Leptasthenura,are in an unresolvedbasal locryptus, rubiginosus, Thripadectes position becausethey build domed or semi- rufobrunneusgroup, and Xenops).Although re- domedvegetative nests inside of cavities(see lationshipswithin cladeA are unresolvedin characters1 and 11). The Schizoeaca-Oreophylax the strict-consensustree, the rootlet-cupOTUs clade (D) is supported by a unique nest of form a clade in some of the fundamental trees Sphagnummoss encapsulatedby herbaceous (eitherimbedded within or sisterto the grass- stems(character 13). The three marsh-nesting cup clade).Among the grass-cupbuilders, the furnariids,Phleocryptes, Limnornis, and Limnoc- cladeof Furnariusis supportedby the clay-oven tites (clade E), are united by constructionof a (character10), and Ochetorhynchuscerthioides is smallawning over the nestentrance (character uniquein lining its nestwith large amountsof 21). Phleocryptesis the only furnariidknown to snakeexuvia (character8). The generaAuto- attach its nest to vertical reed stems (character 900 ZYSKOWSKIAND PRUM [Auk, Vol. 116

22) and build it using plant material coated Our analysissupports a partially resolvedphy- with mud (character12). Clade E composedof logenetichypothesis for the family.Among the Thripophagafusciceps, Siptornis, Premnoplex resulting 15 clades, nest charactersprovide brunnescens,Margarornis, and 11 speciesof the supportfor traditionalas well as novelphylo- Cranioleucaalbiceps group, is diagnosedby pen- geneticgroupings. Although we do not advo- sile nestsof pliable material (character23). In cate the exclusive use of nest characters for contrast,the nestsof the sevenspecies of the C. phylogeneticreconstruction, we present this pyrrhophiagroup are supported from below hypothesisas a heuristictest of utility of nest (i.e. not pensile).Nests of Thripophagafuscicepscharacters in avian systematics.We also use are uniquein havingtwo entrances(character theseresults to proposenew hypotheses for the 19). We failed to find any derivednest charac- evolutionof the diversityof nestarchitecture in tersto diagnoseor resolverelationships among furnariids,which has long been subject to com- Sylviorthorhynchus,Spartonoica, Cranioleuca pyr- parative analyses. rhophiagroup, and Astheneswyatti group. Componentsof variationin nestarchitecture.- Within the stick-nestclade (G), membersof Someof theintraspecific variation in nestplace- Asthenes(except the A. wyattigroup), Acrobator- ment and architectureis adaptivelycorrelated his, Anumbius,Coryphistera, Phacellodomus, and with proximatebiotic and abioticfactors (Col- Eremobiusline not only the bottom area that lias and Collias 1984,Kern and van Riper 1984, cushionsthe eggs,but alsothe walls and ceil- Franklin 1995).As a part of the extendedphe- ing of the incubationchamber (character 15.1). notype,however, nestsare an expression Anumbiusand Coryphisteraform a clade (H) di- of various genotypes in different environ- agnosedby the nest-roofadornment (character ments.Variation in nestphenotype within and 17), and Phacellodomusis supported by the en- amongpopulations can be partitionedinto ge- trance constriction(character 20) and the sec- netic and environmentalcomponents. The ge- ondarilypensile nest (character 24). A nestlin- neticcomponent can evolve and yield fixed dif- ing restrictedto the floor of a brood chamber ferenceswithin lineagesthat may becomein- (character15.2) distinguishesclade I (clade J formative systematiccharacters. The environ- plus Pseudoseisura).Clade J, composedof Cer- mental componentof nest phenotypealso can thiaxis,Schoeniophylax, Gyalophylax, Poecilurus, evolveby selectionfor increasedor decreased and Synallaxis,is supportedby the presenceof plasticity in the face of environmentalvaria- thatch (character16). tion. Comparative studies of nest diversity Relationshipsamong members of cladeG are needto evaluatethe variouscomponents of var- unresolved in the strict-consensus tree. How- iation in nest phenotypeto determinethe rel- ever, a diverse clade of entrance-tube builders ative contributionof historicaland proximate (character18), includingmembers of clade I, factors to the observed variation. Like other Eremobius,Acrobatornis, Phacellodomus, Asthenes systematiccharacters, a behaviormay be phy- humicolagroup, and sometimesthe Asthenes logeneticallyinformative in one lineage but pyrrholeucagroup, appears in many fundamen- variable in another. tal trees with the builders of non-tube stick Sample sizes of nests of severalfurnariids nests as basal to this clade. The entrance-tube were large enoughto permit us to assessmany clade does not appear in the strict-consensus aspectsof intraspecificvariation in nest archi- tree becausethe A. pyrrholeucagroup is poly- tecture.With but few exceptions,furnariids ex- morphicfor characters18 (presenceof entrance hibited low variationwithin speciesand pop- tube), 13 (presenceof sticks),and 1 (nestplace- ulationsfor the nestcharacters hypothesized. ment inside cavity). Even for somepoorly sampledtaxa, limited variationamong species within generaindicat- DISCUSSION ed that phenotypicplasticity within spedes was low. For example,the nestsof eachof the A phylogeneticanalysis of the explosivedi- six speciesof Sclerurusare known from only a versity in nest architecturein the Furnariidae handfulof reports,but eachspecies constructs demonstratesthat the majority of variationin a nest of compound-leafrachises. Other ex- nest constructionis hierarchicallydistributed amples of consistentmaterial preferencesin amonghigher taxa,rather than within species. poorly sampled lineagesinclude use of barn- October1999] NestPhylogeny ofFurnariidae 901 booleaves by Lochmias,tree-fern scales by Pseu- sion, however,was based on only four very docolaptes,and Sphagnummoss by Schizoeaca.general features of swiftletnests. One charac- Suchconsistencies support the conclusionthat ter, the incorporationof feathers,was as phy- variation in furnariid nest-construction behav- logeneticallyconsistent as the averagemolec- ior has been determinedprimarily by natural ular characterused to producethe phylogenet- selectionand less by proximate phenotypic ic hypothesis.Another character, nest proxim- plasticity. ity, actuallyis an aspectof socialbehavior, not Somenest features, however, are quite vari- nest architecture.Detailed analysesof nest ablewithin and amongfurnariids, such as the structure and construction behavior in the Col- useof wool and feathersin the nestlining and localiini and other swifts could yield other the dimensions of nests and cavities. Some in- more-specificand informative nest homologies. traspecificvariation is correlatedwith environ- For example,the use of salivaas an adhesivein mentalvariation in materialavailability. For ex- nest constructionoccurs in all apodimorphs ample,many speciesin cladeG (Fig. 3) seemto (Apodidae + Hemiprocnidae)except Hirunda- prefer thorny sticksfor nest constructionbut pusand Cypseloidinae (Chantler and Driessens use non-thornysticks if the preferredmaterial 1995)and is a clearsynapomorphy of thelarger is unavailable(e.g. Nores and Nores 1994).Ex- clade with a reversal in the latter two taxa. ceptional examplesof extensivegeographic In general,the strengthof a proposedbehav- polymorphismexist in Asthenespyrrholeuca and ioral character is related to the amount of detail A. modesta.Various populations construct supportingthe primaryhypothesis of homol- domednests of thorny stickswith tubular en- ogy (sensuPinna 1991). Unlike many behav- trancesin the branchesof shrubs,whereas oth- iors, nest construction is amenable to detailed er populationsconstruct reduced, cup-shaped analysisbecause nests can be collected,dis- nests without sticks in burrows or other cavi- sected, and examined thoroughly (Hansell ties. No known geographicor ecologicalgra- 1984, Collias 1986). Furthermore, nest-con- dients are correlatedwith this variation(Con- structionbehaviors can be quite elaborate.For treras1980). Further study is requiredto elu- example,Thaler (1976) distinguished 15 behav- cidate the causesof this polymorphism,but ioral actsperformed during nest construction multiple speciesmay actuallybe involved,es- by kinglets(Regulus spp.). Although such be- peciallyin the polytypicA. modesta. haviorsare poorly known in furnariids,some Phylogeneticutility of nestcharacters.--The co- speciesemploy several distinct stereotyped herent patternsof nest homologyshared by movementsduring nest construction(Vaz-Fer- furnariidlineages indicate that nestplacement, reiraet al. 1993).Detailed analyses of the mod- structure,and constructionbehavior can pro- al actionpatterns involved in nestconstruction vide informativephylogenetic characters. The couldprovide a richsource of phylogeneticin- ultimatetest of the phylogeneticutility of nest formation.In addition,the ontogenyof nests charactersin furnariidswill be providedby can bring specialdetail to hypothesesof ho- congruencewith independentlyderived data mologyand providerobust phylogenetic char- sets. acters(see below). Architectural characters are A few studieshave successfullyused nest phylogeneticallyinformative in severalgroups charactersin phylogeneticreconstruction (e.g. of non-avian speciesas well (Hansell 1984, Lanyon 1986, 1988). Recent analysesof avian Wenzel1992). This resultfurther supportsthe nest features have reached diverse conclusions utility of nestcharacters in phylogeneticrecon- aboutthe phylogeneticcontent of nest varia- struction. tion. For example, Prum (1993), Winkler and Evolutionof nest architecturein furnariids.-- Sheldon(1993), Nunn et al. (1996), Eberhard Whethermaintained by naturalselection or by (1998), and Sturmbaueret al. (1998) document- phylogeneticconstraint, enclosed nests occur ed substantialcongruence between nest archi- throughoutthe Furnariidaeand are achieved tecture and phylogeny,indicating that nests by a varietyof architecturalsolutions in mul- can provide valuablesystematic characters. In tiple lineages(Fig. 4). As Skutch(1996:161) contrast,Lee et al. (1996) concludedthat nest stated, "the only feature that all [furnariid characterswere not phylogeneticallyinforma- nests]have in commonis their provisionof a tive within swiftlets(Collocaliini). This conclu- domed,roofed spacefor eggs,nestlings, and 902 ZYSKOWSKIAND PRUM [Auk, Vol. 116 attendantparents .... "Comparisonswith the rion, featuresappearing earlier in ontogenyare outgroupsdemonstrate that cavitynesting is morebroadly distributed among taxa, whereas plesiomorphicto thefamily. In thetwo lineages featuresacquired in subsequentdevelopmental wherethe primitivecavity nest has been lost, stageshave narrower distributions and arehy- novel structures have evolved to enclose the pothesizedas derived.The ontogeneticcrite- nest contents:the clay oven of Furnarius,and rion performswell when noveltiesresult from the domedvegetative nest of the synallaxine terminaladditions to developmentalsequences clade. Even the simple nests of Spartonoica,(Meier 1997). which someauthors interpret as the lossof a The ontogeneticcriterion can often be used dome (Collias1997), essentially are enclosedin to determinepolarity of nest charactersbe- thickvegetation that providesa completecov- causeof thephysical or engineeringconstraints eringfor the nest. that are imposedon the developmentof three- Our phylogenysuggests several specific hy- dimensional constructions.For example, be- pothesesabout the historyof nestevolution in cause no avian nests are constructed and sub- the Furnariidae.For example,the domedclay sequentlyattached to a substrate,the necessity nestof Furnariusis uniquein birds.Doello-Ju- of attachmentor placementconstrains the on- rado (1919)proposed that the clayoven of Fur- togenyand evolutionof nests.The primitive nariusis homologousto the domed nest of form of the domed stick nest found in clade G mud-daubed plant material in Phleocryptes.is a platformsupported from below.The sec- However, our phylogenetichypothesis indi- ondarily pensilestick nest of Phacellodomus,in catesthat the discretegrassy cup in the adobe whichthe nestis initially supportedfrom be- dome made by Furnariusis not homologous low but becomespensile as its increasingmass with the vegetativenest of Phleocryptes.Rather, causesthe supportingbranch to collapse,is our resultsindicate that the claydome of Fur- uniqueamong birds. The primitiveontogeny nariusis homologousto the burrowor cavities within thisgroup (i.e. platform supported from of otherfurnariids, and that the grasscup is a below) precludedthe evolutionof a typical homologshared with othercavity-nesting fur- pensilenest, which is built from the top down, nariids.This hypothesisis further supported but this constraint also fostered the evolution of by the observationthat someFurnarius are fac- a truly novelmethod of constructinga hanging ultative or obligatecavity nesters (Sclater and domed nest. Salvin 1873, Studer and Vielliard 1990, Sick Winklerand Sheldon(1993) used nest ontog- 1993,K. Zyskowskipers. obs.). eny to polarizevariation in nest architecture Nestontogeny.--In addition to the nest and within mud-nestingswallows (Hirundinidae). the behavioralacts used to constructit, a great They argued that closed and retort nests dealof informationexists in theontogeny of the evolvedfrom an open mud cup becausethe nest.The sequence of eventsin theconstruction nest beginsas a cup that is subsequentlyen- of nests may reveal additional information closed, and to which the retort entrance tunnel aboutthe hornologyof crypticallyhomologous is finally added.It is likely that the ontogenetic or superficiallysimilar but analogousnests. For criterionwill have other successfulapplica- example,the nestof mostPhacellodomus begins tionsto polarizationof avian nestcharacters. as a mat of sticksnear the tip of a horizontal Phylogenyof the Furnariidae.--Ouranalysis branchand eventuallybecomes pensile as the supportsa partiallyresolved phylogenetic hy- supportingbranch droops under the massof pothesisfor the Furnariidae.The phylogenyin- thenest. In contrast,pensfie nests of Thripopha- cludesthree novel basal clades(clades A to C) gafusciceps (placed in Phacellodomusby Vaurie and three genera (Chilia, Pygarrhichas,and [1980])are built by draping strandsof vegeta- Pseudocolaptes)with unresolvedrelationships. tionfrom the supportingbranch downward. In The largestresolved group (clade C) includes thisinstance, a superficial similarity that could 27 generathat build domedvegetative nests. be a proposedhornology, "pendant nest," is This nest type unitesnot only all traditional falsifiedby ontogeneticinformation. Synallaxinaeanalyzed but also Eremobius(tra- Ontogenyhas alsobeen proposed as a cri- ditionallyplaced in Furnariinae)and Margaror- terionfor polarizingcharacter variation among nis, Premnoplex,Pseudoseisura, and Lochmias taxa (e.g.Meier 1997).According to this crite- (usuallyplaced in Philydorinae).Within this October1999] NestPhylogeny ofFurnariidae 903 expanded"synallaxine" clade,additional nest ditionalevidence is required to confirmthis no- synapomorphiesprovide supportfor several tion. more-resolvedsubclades (D to J), including severalnovel phylogeneticrelationships. For ACKNOWLEDGMENTS example,we found behavioralsupport for a clade (E) that includesthe three monotypic We are extremelygrateful to the followingfor pro- marsh-nestinggenera (Limnornis, Limnoctites, viding nestand egg data, lendingphotographs and nest specimens,and assistingduring visits to their and Phleocryptes)and for a clade(J) thatunites collections: M. LeCroy (AMNH), M. Walters Certhiaxiswith the Synallaxisassemblage (sen- (BMNH), G. Hess (DMNH), J. Batesand D. Willard su Vaurie 1980). (FMNH), T. Stjernberg (HMNH), J. V. Remsen We alsoidentified a nestsynapomorphy to (LSUMZ), D. Teixeira and J. Nacinovic (MNRJ), D. supporta clade(F) thatincludes the Cranioleuca Oren (MPEG), I. Franke (PMHN), R. McKernan albicepsgroup, Margarornis, Premnoplex brunnes- (SBCM),J. Dean and G. Graves(USNM), B. Mathews cens,Siptornis, and Thripophagafusciceps. Al- and D. Wechsler(VIREO), and S. Sumida and E Ki- thougha relationshipof Siptornisto Cranioleucanoshita (WFVZ). We also thank the following for sharingunpublished descriptions of furnariid nests: has been proposedpreviously (Vaurie 1980), T. Abreu, E. Barnes,A. Begazo,C. Brady,C. Byers,D. thisentire assemblage isnovel. Until now,these Christian, M. Cohn-Haft, N. Collias, T. Davis, J. generahave been placedin two subfamilies Fjelds•, P. Greenfield,S. Hilty, A. Jaramillo,L. Kiff, (Cranioleucaand Thripophagain Synallaxinae; D. Lane, M. Marin, P. Martin, R. McNeal, T. McNish, Margarornisand Premnoplexin Philydorinae), E. Mendonqa,J. Minns, J. O'Neill, R. Parrini, M. Ra- despitethe apparentbehavioral and vocalsim- poso,P. Rasmussen,M. Reid, M. Robbins,D. Robin- ilaritiesamong them (Ridgely and Tudor 1994). son,T. Ryan,J. Sfinchez,T. Schulenberg,C. Sharpe, G. Stiles,B. Walker,D. Watson,N. Wheelwright,B. The monophylyof two diversegenera (As- Whitney,and A. Whittaker.For assistancein the field thenesand Cranioleuca)is not supportedby our and permissionto collectnests, KZ is indebtedto results.In the caseof Asthenes,the A. wyatti employeesof the Museo Nacional de Historia Natu- groupis a basalmember of the domedvege- ral del Paraguay,CITES, Fundaci6n Moises Bertoni, tativenest clade (C), whereasthe A. pyrrholeucaEstancia Zalazar, participantsof the Yacutinga'95 and A. humicolagroups, and A. anthoides,are Project (University of Cambridge), and H. Amarilla, membersof clade G, which is diagnosedby M. Rfiberg,and M. Robbinsin Paraguay;employees highly derivedstick nests. These "grass-nest- of Instituto Nacional de Recursos Naturales, Manu Expeditions, Selva Sur, Amazonia and Pantiacolla ing" and "stick-nesting"assemblages of Asthe- lodges,and H. Amarilla in Peru;and R. Macedo,E. neshave been recognized previously, and they Mendonqa,J. Minns, D. Oren, R. Parrini, M. Raposo, can also be diagnosedusing plumage charac- and managersof Casa da Bocainaand Borac•ia Bi- ters and vocalizations(Narosky et al. 1983,Pa- ologicalStation in Brazil.We alsothank J. Eberhard, checoet al. 1996).In contrast,the two groups K. McCracken,R. Zink, and an anonymousreviewer of Cranioleucarecognized here based on nest ar- for improving the manuscript.This researchwas chitecturedo not parallelthose identified using supportedby grantsfrom J. Zyskowski, the Frank M. molecular,morphological or distributionalev- ChapmanMemorial Fund (AMNH), GraduateStu- dent Scholarship(DMNH), PanoramaSociety, Goss idence(Maijer and Fjeldsg1997, J. Garc•a-Mo- Fund, and Tinker Field ResearchGrant (University reno pers. comm.). of Kansas)to KZ, andby NSF grantDEB-9318273 to Relationshipsamong the cavity-nestinglin- ROP. eages of fumariids are not so well resolved. However,these lineages include several clades LITERATURE CITED diagnosedby distinctivenest synapomorphies. The largest of these groupsis the cup-nest ALLEN,J. A. 1905.Supplementary notes on birds col- clade (A) that includesFurnarius. Although lectedin the SantaMarta District, ,by Furnariustraditionally has been associated Herbert Smith, with descriptionsof nests and with manyof the generain cladeA, our study eggs.Bulletin of the AmericanMuseum of Nat- ural History 21:275-295. has identifiedthe first behavioralsynapomor- AMES,P. L. 1971. The morphologyof the in phy that supportsthis relationship.Our phy- passerinebirds. Bulletinof the PeabodyMuse- logenetichypothesis also suggeststhat Auto- um of Natural History 37:1-194. molusand Thripadectesare polyphyletic, but ad- ANDRADE,M. A. DE. 1996. Observaq•essobre ninhos 904 ZYSKOWSKIAND PRUM [Auk, Vol. 116

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Parsimony, version 3.1.1. Computer program ciones del Museo de Historia Natural de distributedby the author,Smithsonian Institu- UNMSM, SerieZoo16gica (A) 32:1-7. tion, Washington,D.C. VUILLEUMIER,F. 1968. Population structure of theAs- TACZANOWSKI,L. 1884. Ornithologiedu P•rou, vol. thenesfiarnrnulata superspecies (Aves, Furnari- 2. Friedl•inder und Sohn, Berlin. idae). Breviora 297:1-21. TEIXEIRA,D. M. 1992.Sobre a biologiada "Maria-Ma- VUILLEUMIER, E 1969. Field notes on some birds from cambira," Gyalophylaxhelltnayri (Reiser, 1905) the Bolivian Andes. Ibis 111:599-608. (Aves,Furnariidae). Iheringia (S•rie Zoo16gica) VUILLEUMIER,E 1993.Field study of allopatry,sym- 72:141-146. patry, parapatry, and reproductive isolation in TEIXEIRA, D. g., AND G. LUIGI. 1989. Notas sobre steppebirds of Patagonia.Ornitologla Neotrop- Cranioleuca semicinerea (Reichenbach, 1853) ical 4:1-41. (Aves, Furnariidae). Revista Brasileira de Biolo- WAGNER,H. O. 1980.Vogelleben im mexikanischen gia 49:605-613. Regenwald.Gefiederte Welt 104:95-97. TEIXEIRA, D. g., AND G. LUIGI. 1993. Notas sobre Poe- WALTERS,R. 1995.Notes on the eggsof the Scaly- cilurus scutatus(Sclater, 1859) (Aves, Furnari- throatedLeaftosser, Sclerurus guatetnalensis. Or- idae). Iheringia(S•rie Zoo16gica)74:117-124. nitologia Neotropical6:53. THALER, E. 1976. Nest und Nestbau von Winter- und WENZEL,J. W. 1992.Behavioral hornology and phy- logeny.Annual Reviewof Ecologyand System- Sommergoldh•ihnchen(Regulus regulus und R. atics 23:361-381. ignicapillus).Journal ftir Ornithologie117:121- WHITNEY, B. M., J. E PACHECO,P.S. M. DA FONSECA, 144. AND R. H. BARTH.1996. The nest and nesting THOMAS, B. T. 1983. The Plain-fronted Thornbird: ecology of Acrobatornisfonsecai (Furnariidae), Nest construction, material choice,and nest de- with implicationsfor intrafamilialrelationships. fense behavior Wilson Bulletin 95:106-117. Wilson Bulletin 108:434-448. TOSTAIN, O., J. L. DUJARDIN, C. ERARD, AND J. M. W•LLIAMS,M.D. 1981. The nest and eggs of the THIOLLAY.1992. Oiseaux de Guyane. Soci•t• CoastalMiner GeosittaperuEiana, a Peruvianen- d'EtudesOrnitologiques, Brunoy, France. demic. Ibis 123:366-367. VAur•m,C. 1971.Classification of the ovenbirds(Fur- W•LLI$,E. O. 1988.Behavioral notes, breeding rec- nariidae).H. E & G. Witherby,London. ords, and rangeextensions for Colombianbirds. VAur•m,C. 1980. and geographicaldistri- Revista de la Academia Colombiana de Ciencias bution of the Furnariidae (Aves,Passeriformes). ExactasFisicas y Naturales16:137-150. Bulletin of the American Museum of Natural WINKLER,D. W., AND F. H. SHELDON.1993. Evolution History 166:1-357. of nest constructionin swallows (Hirundinidae): VAZ-FERREIRA,R., E. PALERM,J. BIANCO,AND A. STA- A molecularphylogenetic perspective. Proceed- GI. 1993. Caracterlsticasdel nido y t•cnicas de ings of the National Academyof SciencesUSA construcci6ndel "hornero"Furnarius rufus y del 90:5705-5707. "espinero" Anutnbiusannutnbi (Furnariidae, Pas- Z•MMER,K. J. 1997. Specieslimits in Cranioleucavul- seriformes).Boletin de la SociedadZoologica del pina.Pages 849-864 in Studiesin Neotropicalor- Uruguay 8:239-264. nithologyhonoring Ted Parker(J. V. Reinsen,Jr., VENERO,J. L. 1990. Clasificaci6n de nidos de aves (en Ed.). OrnithologicalMonographs No. 48. base a especiesde la puna del Per6). Publica- Associate Editor: R. M. Zink 908 ZYSKOWSK!AND PRUM [Auk, Vol. 116

APPENDIX1. Sources of information on nest architecture of furnariids. Species are arranged in alphabetical order with generic names in bold. References are listed separately for subspecifictaxa that have been hypothesizedto deserve speciesrank (Ridgely and Tudor 1994). All suspectedcases of uncertain species identification are coded "id." Rather than providing an exhaustive list of references from the scientific literature, we selected a minimal set of publications for each species,giving priority to compilations and more recent papers. Published information is followed by unpublishedphotographic or specimen-based sources. Specimen data and VIREO images are listed as collector's or photographer'sname followed by institution code and catalog number(s). Information obtained through personal communicationsis presented by listing a contributor'sname alone. Abbreviationsused for frequently cited sourcesare: B84 (Belton 1984), FK90 (Fjeldsfi and Krabbe 1990), HB86 (Hilty and Brown 1986), J67 (Johnson1967), N83 (Narosky et al. 1983), P87 (de la Pefia 1987), S93 (Sick 1993), V80 (Vaurie 1980), AW (A. Whittaker), BW (B. Whitney), and KZ (K. Zyskowski). Institutional codes are: AMNH (American Museum of Natural History, New York), BMNH (Natural History Museum, Tring), DMNH (Delaware Museum of Natural History, Greenville), FMNH (Field Museum of Natural History, Chicago), HMNH (Finnish Museum of Natural History, Helsinki), KU (University of Kansas Natural History Museum), MNRJ (Museu Nacional, Univ- ersidade Federal de Rio de Janeiro), MPEG (Museu ParaenseEmilio Goeldi, Belfim, Brazil), PMHN (Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima), SBCM (San Bernardino County Museum, Redlands, California), USNM (National Museum of Natural History, Smithsonian Institution), WFVZ (Western Foundation of Vertebrate Zoology, Camarillo, California), and VIREO (Visual Resources for Ornithology, Academy of Natural Sciences of Philadelphia).

Acrobatornis fonsecai: Whitney et al. 1996; striaticollis: V80 (Schaefer, HMNH 13249), T. McNish (VIREO M42/1/092-094, pers. comm.); A. variegaticeps: Wagner 1980; Anabazenops dorsalis: Kratter 1994; A. fuscus: S93, BW; Anumbius annumbi: V80, N83, B84, Mason 1985, P87, S93, Vaz-Ferreira et al. 1993, Belcher (HMNH 13079), Crossin (WFVZ 160674-7), Holt (USNM 41089), MacIntyre (DMNH 23685), Ridgely (VIREO R10/13/064-8), Young (DMNH 23684), R. Parrini, AW, KZ; Aphrastura spinicauda: J67 (Johnson and Goodall, WFVZ 55514-9), V80, N83, P87, Crossin (WFVZ 160700), P. Rasmussen (VIREO R17/1/017, pers. comm.); Asthenes anthoides:N83, Colias 1997, Belts (DMNH uncat., id); A. arequipae: J67 (Johnson and Goodall, WFVZ 55551), V80, FK90, Dorst (WFVZ 151796); A. baeri: N83, B84, P87, Girard (WFVZ 53772), KZ; A. berlepschi:Collar et al. 1992, BW; A. cactorum:Koepcke 1959, V80, T Schulenberg; A. dorbignyi: V80, N83, Fraga and Narosky 1985, P87; A. huancavelicae:Morrison 1939, FK90, Collar et al. 1992, Hocking (PMHN uncat.); A. hudsoni: V80, N83, Goodall (DMNH 23671), Wetmore (USNM 34128), Young (DMNH 23672); A. humicola: J67 (Goodall and Johnson: DMNH 23664-70; USNM 39973; WFVZ 23408, 53168, 55540-50), N83, P87, Marin (WFVZ 120289); A. humills: V80; A. luizae: BW; A. maculicauda: KZ; A. modesta: J67 (Goodall and Johnson: WFVZ 53169-70, 55552-65), V80, N83, P87, FK90, Venero 1990, Pemberton (WFVZ 15148-58), BW; A. patagonica: V80, N83, Fraga and Narosky 1985, FK90, Vuil- leumier 1993, Whitney et al. 1996, Wetmore (USNM 36250); A. pudibunda: Koepcke 1958; A. pyrrholeuca: J67 (Goodall and Johnson: WFVZ 55636-7), V80, N83, P87, Pemberton (WFVZ 15144-7), Smyth (DMNH uncat., HMNH 13060), Wetmore (USNM 36249); A. sclateri: N83, Fraga and Narosky 1985, P87; A. steinbachi: N83, Salvador 1992; A. urubambensis:KZ; A. virgata: BW; A. wyatti aequatorialis group: HB86, Phelps 1977; A. w. graminicola: Dorst 1963 (WFVZ 151795), BW; Automolus infuscatus:Pinto 1953, Koepcke 1972, V80, Tostain et al. 1992; A. leucophthalrnus:V80, N83, M. Raposo, AW, KZ; A. melanopezus:E. Barnes; A. ochrolaemus: Skutch 1969, V80, Kiff (WFVZ 58600-1), Stiles (WFVZ 58620), Williams (WFVZ 59367); A. rubiginosus:Rowley 1966 (WFVZ 21315), V80, Marfn and Carrion 1991 (WFVZ 158304); Berlepschia rikeri: V80, S93; Certhiaxis cinnamomea: Cherrie 1916 (WFVZ 120131), Belcher and Smooker 1936 (DMNH 23662; HMNH 11976-8, 11980, 13056-7), V80, N83, B84, HB86, P87, S93, Haverschmidt and Mees 1994, Carriker (WFVZ 154606, 154984; SBCM 20184-92, 20383, 20433), Contino (WFVZ 156061, 158445), Crandall (DMNH 23661, WFVZ 117042), MPEG uncat., KZ; C. mustelina: V80, BW; Chilia melanura: J67 [Behn (WFVZ 55493), Goodall and Johnson (USNM 39975; WFVZ 53175-6, 55489-92)], V80, M. Marfn; Cinciodes antarcticus: Pettingill 1973, V80, N83, Belts (DMNH, 3 uncat.); C. atacamensis: Morrison 1939, J67 (WFVZ 55508), N83, Salvador and Narosky 1984, P87, Ralde (WFVZ 55509); C. comechingonus: N83, P87; C. excelsior: Graves and Arango 1988, Marin et al. (WFVZ 162880-1); C. fuscus: J67 (Johnsonand Goodall: WFVZ 53114-5, 55504-7), V80, N83, P87, Venero 1990, Salvador 1992, Carrion (WFVZ 161930), Kiff et al. (WFVZ 162874), Pemberton (WFVZ 15132-4), KZ; C. nigrofumosus:J67 (Goodall, WFVZ 55503; Millie, WFVZ 55502), V80; C. olrogi: N83, P87; C. oustaleti: J67 (Goodall, WFVZ 55510), N83; C. pabsti: B84, S93; C. patagonicus: J67 (Goodall: WFVZ 53110-3, 55494-501), N83, Pemberton (WFVZ 15135); C. tac- zanowskii: V80, BW; Coryphistera alaudina: V80, N83, P87, Dinelli (WFVZ 150668), Girard (WFVZ 53759), R. Parrini, KZ; Cranioleuca albicapilla: Peters and Griswold 1943, Parker and O'Neill 1980 (id), E. Barnes, J. Fjeldsfi; C. albiceps: Remsen 1984; C. antisiensis:Koepcke 1958, D. Lane; C. baroni: FK90; C. curtata: M. Cohn-Haft, G. Stiles; C. demissa: BW (id); C. erythrops: Skutch 1969, V80, Willis 1988, Skutch 1996, Marfn (WFVZ 157422), G. Stiles (WFVZ 68520, pers. comm.); C. gutturata: HB86, E. Barnes, M. Reid, M. Robbins, AW, KZ; C. hellmayri: HB86; C. marcapatae: Parker and O'Neill 1980 (id), KZ; C. muelleri: BW, AW; C. obsoleta: B84; C. pallida: V80, S93, AW, BW, KZ; C. pyrrhophia: V80, N83, B84, P87, Girard (WFVZ 53771), R. Parrini; C. semicinerea: Teixeira and Luigi 1989; C. subcristata: V80; C. sulphurifera: V80, N83, P87, Runnacles (SBCM 22271); C. vulpina: Cherrie 1916 (WFVZ 120131), Teixeira and Luigi October 1999] Nest Phylogenyof Furnariidae 9O9

APPENDIX 1. Continued.

1989, Holt (USNM 41091-2); Eremobius phoenicurus: V80, N83, Vuilleumier 1993, Crossin (WFVZ 160701), Pemberton (WFVZ 15136-40); Furnarius cristatus: N83, P87, KZ; F. figulus: V80, Studer and Vielliard 1990, S93, AW, KZ; F. I. cinnamomeus: Taczanowski 1884, Marin in Skutch 1996, Kiff et al. (WFVZ 162164), D. Lane, KZ; F. I. leucopusgroup: Ihering 1914, Naumburg 1930, Koepcke 1972, Dyrcz 1987, S93, Smooker (HMNH 13070-1), KZ; F. I. Iongirostris: Allen 1905, Carriker (SBCM 20356, 20358); F. minor: Sclater and Salvin 1873, Chagas (WFVZ 24442); F. rufus: V80, N83, B84, Mason 1985, P87, S93, Vaz-Ferreira et al. 1993, Blake (FMNH 2269, 14756), Chagas (WFVZ 24443, 53777), Crossin (WFVZ 160680-3), Dunlap (USNM 40707), Garrett (DMNH 23633), Holt (USNM 41090), Mason (WFVZ 146877), Meyer (USNM 47248), Sobrinho (HMNH 13067), Wetmore (USNM 36251, 36333), Young (DMNH 23632), MNRJ uncat., KZ; F. torridus: Sclater and Salvin 1873; Geobatespoecilopterus: S93; Geositta antarctica: J67 (Johnsonand Goodall, WFVZ 55478), N83, Hermiston (DMNH uncat.); G. cunicularia cunicularia group: J67 (Johnsonand Goodall: WFVZ 53103-4, 55470-4), N83, B84, P87, S93, Belcher (HMNH 13064), Marin (WFVZ 129185), Pemberton (WFVZ 15128-9); G. c. deserticolor: J67 (Millie, WFVZ 55475-6); G. c. frobeni group: J67 (Johnson and Goodall, WFVZ 55477), Venero 1990, R. McNeal; G. isabellina: V80, N83; G. maritima: J67; G. peruviana: Williams 1981, D. Lane, KZ; G. punensis: J67, V80, N83, Ralde (WFVZ 55481); G. rufipennis: J67 (Johnson and Goodall, WFVZ 55479-80), V80, N83, P87, Pemberton (WFVZ 15130); G. saxicolina: Taczanowski 1884; G. tenuirostris: Taczanowski 1884, Salvador et al. 1984, R. McNeal; Gyalophylax hellmayri: Ihering 1914, Teixeira 1992, S93, AW; Hylocryptus erythrocephalus: Collar et al. 1992, M. Robbins, KZ; H. rectirostris: S93, T Abreu, R. Parrini; Hyloctistes s. subulatus: Sclater and Salvin 1873 (id); H. s. virgatus: N. Wheelwright; Leptasthenura a. aegithaloides: J67 (Goodall and Johnson:WFVZ 53031-7, 55532-6); L. a. berlepschi: J67 (Johnson and Goodall, WFVZ 55539), N83, P87; L. a. grisescens:J67 (Millie, WFVZ 55538); L. a. pallida: N83, Fraga and Narosky 1985, P87, Vuilleumier 1993, Pemberton(WFVZ 15141-3); L. andicola: V80, HB86, Briceno (USNM 30597), Fjelds• and Krabbe(WFVZ 162898), Kiff andGarrett (WFVZ 162947), Marin (WFVZ 162891), M. Robbins (VIREO R08/9/024, pers. comm.), C. Sharpe;L. fuliginiceps: V80, N83, Contino (WFVZ 156116); L. pileata: I. Franke (PMHN uncat., pers. comm.); L. platensis: V80, N83, B84, Mason 1985, P87, S93, MacIntyre (DMNH uncat.), Malcolm (DMNH 23634), Short (WFVZ 160691); L. setaria: B84, Andrade 1996, AW; L. striolata: B84, BW; L. xenothorax: C. Byers and KZ; L. yanacensis: Vuilleumier 1969; Lim- noctites rectirostris: N83, Ricci and Ricci 1985, S93; Limnornis curvirostris: V80, N83, B84; Lochmias nematura: V80, N83, B84, S93, AMNH 13846-7, T. Abreu, T Davis, BW, KZ; Margarornis rubiginosus: J. S•nchez, D. Watson; M. squamiger: HB86, FK90; Metopothrix aurantiacus: Fraga 1992, Ridgely and Tudor 1994, A. Begazo, P. Greenfield, R. McNeal, M. Robbins, B. Walker, BW; Ochetorhynchus certhioides: N83, P87, KZ; Oreophylax moreirae: Sick 1970, S93; Phace!!odornus dorsalis: Ridgely and Tudor 1994, J. O'Neill (VIREO O03/2/178, pets. comm.), B. Walker; P. e. erythrophthalmus: Euler 1900, S93, E. Mendonqa, KZ; P. e. ferrugineigula: Ihering 1900, V80, R. Parrini; P. maculipectus: N83, Salvador 1992, Mazar Barnett et al. 1998; P. ruber: V80, N83, S93, P87, Crossin (WFVZ 160690), Pettingill (VIREO P03/7/075), AW, KZ; P. ruffirons inornatus group: Skutch 1969, Thomas 1983, HB86, Phelps and Aveledo 1987, Collias 1997, C. Brady; P. r. peruvianus: Taczanowski 1884, J. O'Neill (VIREO O03/2/177), D. Lane, KZ; P. r. ruffirons group: Naumburg 1930, N83, P87, S93, Whitney et al. 1996, Girard (WFVZ 53774), AW, KZ; P. sibilatrix: V80, N83, Mason 1985, P87, KZ; P. striaticeps: V80, N83, P87, KZ; P. striaticollis: V80, N83, B84, Mason 1985, P87, S93; Philydor atricapillus: Ruschi 1981 (id), S93, BW, KZ; P. erythrocercus: Tostain et al. 1992; P. lichtensteini: Saibene 1995, AW, BW; P. rufus columbianusgroup: V80 (Schaefer, HMNH 13254; Schwartz in "A Portfolio of Venezuelan Birds," undated publ. of Cornell Lab. Orn.); P. r. riveti: HB86; P. r. rufus: V80, Ruschi 1981 (id), N83, BW, J. Minns, KZ; Phleocryptes melanops: Taczanowski 1884, J67 (Goodall and Johnson: DMNH 23677; USNM 39977; WFVZ 53021-30, 55520-31), V80, N83, B84, Mason 1985, P87, S93, Felippone (USNM 37417-8), Garrett (DMNH 23679), MacIntyre (DMNH 23678), Marin (WFVZ 120291, 129189-92), Pefia (USNM 2730), Wetmore (USNM 34127) Williams (WFVZ, 3 uncat.), KZ; Poecilurus candei: V80, Bosque and Lentino 1987; P. scutatus: Teixeira and Luigi 1993, SBCM 20362; Premnoplex brunnescens:Skutch 1967 (WFVZ 158848), V80, FK90, Marin and Carrion 1994 (WFVZ 157401, 159225, 162250, 162895-6), Sibley (WFVZ 157319, labeled as Lochrniasnernatura); Pseudocolaptes boissonneautii: Sclater and Salvin 1879; P. lawrencii: Skutch 1969, G. Stiles (WFVZ 151475, pets. comm.); Pseudoseisura cristata: Ihering 1914, Naumburg 1930, V80, S93, AW; P. gutturalis: V80, N83, P87, Collias 1997, Pemberton (WFVZ 15159-60); P. Iophotes: V80, N83, P87, S93, Notes and Nores 1994, Goodall (DMNH 23690), Maclntyre (DMNH 23686), Malcolm (DMNH 23689), Smyth (HMNH 13074), KZ; Pygarrhichas albogularis: J67 (Goodall and Johnson(WFVZ 55567-9), V80, N83, FK90; $chizoeaca fuliginosa: HB86, M. Robbins; S. griseomurina:D. Christian; S. harterti: Vuilleumier 1969; S. helleri: T. Schulenberg(WFVZ, 2 uncat.; pets. comm.); Schoeniophylax phryganophila: V80, N83, B84, P87, S93, Crossin (WFVZ 160678-9), MacIntyre (DMNH uncat.), AW, KZ; Sclerurus albigularis: Belcher and Smooker 1936 (HMNH 13061-3), V80, HB86, Carriker (WFVZ 154607-8; SBCM 20113-22, 20193-8), T. Davis; S. caudacutus: Pinto 1953, KZ; S. gua- temalensis: Skutch 1969, V80, Walters 1995, D. Robinson, Stiles (WFVZ 64490), Williams (WFVZ 58180); S. mexicanus: Tostain et al. 1992, Stiles (WFVZ 143838, 158873); S. rufigularis: Haverschmidt and Mees 1994, M. Cohn-Haft; S. scansor: V80, N83, S93, Sobrinho (DMNH 23691), AW, M. Raposo, KZ; $iptornis striaticollis: Ridgely and Tudor 1994, P. Greenfield; $iptornopsis hypochondriacus:Braun and Parker 1985, T. Schulenberg,M. Robbins; Spartonoica maluroides: V80, N83; Sylviorthorhynchus desmursii:J67 910 ZYSKOWSKI AND PRUM [Auk, Vol. 116

APPENDIX 1. Continued.

(Goodall, WFVZ 55513), V80, N83; Synallaxis albescens:Sclater and Salvin 1879, Cherrie 1916 (WFVZ 119328, 120127-8), Belcher and Smooker 1936 (DMNH 23655; HMNH 11662-6, 13080), Skutch 1969, V80, N83, P87, Tostain et al. 1992, Hayerschmidt and Mees 1994, Fiala (WFVZ 68051), Smyth (DMNH 23640), Stiles (WFVZ 141906), Young (DMNH 23639), KZ; S. albigularis: V80, HB86; S. albilora: Naumburg 1930 (photos, id); S. a. azarae group: Taczanowski 1884, KZ; S. a. elegantior group: Taczanowski 1884, HB86, KZ; S. a. superciliosa: N83, P. Martin; S. brachyura: Skutch 1969, V80, HB86, Kiff (WFVZ 58437, 58466, 58583-5, 78161, 158324, 158336, 158340), Smith (USNM 39797, WFVZ 69290-2), Stiles (WFVZ 58626- 7), Williams (WFVZ 58441); & chinchipensis: KZ; S. cinerascens: B84, Simon and Pacheco 1996; S. cin- narnomea carri: Belcher and Smooker 1936 (HMNH 11671-2, 11979), V80; S.c. cinnarnomea group: V80, HB86; S.c. terrestris: Belcher and Smooker 1936; S. erythrothorax: Lea 1951, Skutch 1969 and 1996, V80, Skinner (USNM 32475-9); S. frontalis: V80, N83, P87, Girard (WFVZ 53760), KZ; S. gujanensis: Snethlage 1935, Pinto 1953, HB86, Hayerschmidt and Mees 1994, Myers (HMNH 13081), G. Stiles, KZ; S. rnacconnelli: AW; S. propinqua: BW; S. ruficapilla: V80, N83, S93, Chagas (WFVZ 24485; SBCM 20880-1), KZ; S. rutilans omissa: Pinto 1953, Chagas (WFVZ 24487); & r. rutilans: Hayerschmidt and Mees 1994; S. spixi: V80, N83, B84, Fraga and Narosky 1985, P87, Isoldi 1992, S93, Chagas (WFVZ 24486), Smyth (DMNH 23656, 23660), Sobrinho (DMNH 23657-9), E. Mendonqa, KZ; S. stictothorax: Taczanowski 1884, Marchant 1960, Corado and Kiff (WFVZ 158327-8, 158330, 158373), M. Mar/n, KZ; S. subpudica: HB86; S. tithys: Balchin 1996, KZ; S. zimrneri: I. Franke (PMHN uncat., pers. comm.); guttulata: V80 (id); & rufosuperciliata: N83, B84, KZ; Thripadectesfiammulatus:V80, HB86, G. Stiles; T. holostictus:P. Alden (VIREO A01/1/371, pers. comm.; id), BW; T. rnelanorhynchus:Kiff et al. 1989 (WFVZ 156103); T. rufobrunneus:Skutch 1969, Marfn and Schmitt (WFVZ 154831), G. Stiles, Worth (WFVZ 154898); T. scrutator: M. Robbins, BW; T. virgaticeps: HB86, Marin and Carrion 1994 (WFVZ 159222); fusciceps: Whitney et al. 1996, E. Barnes, S. Hilty, M. Cohn-Haft, AW, KZ; T. macroura: Collar et al. 1992, Whitney et al. 1996, R. Parrini, AW; Upucerthia albigula: J67 (Johnsonand Goodall, WFVZ 55487), Schulenberg 1987, FK90; U. andaecola: N83, Fraga and Narosky 1985, P87; Uodurnetaria: J67 [Goodall and Johnson(DMNH, 2 uncat.; WFVZ 53171- 3, 55482-4), Millie (WFVZ 55485-6)], V80, N83, P87, Pemberton (WFVZ 15131); U. jelskii: Taczanowski 1884, N83, Venero 1990, Mazar Barnett et al. 1998; U. ruficauda: J67, N83, P87; U. serfaria: FK90, V80; U. validirostris: N83, P87, Salvador 1992; rninlosi: Ridgely and Gwynne 1989 (id), Whitney et al. 1996 (id), M. Reid; X. singularis: T Davis (id), P. Greenfield; Xenops rnilleri: AW; X. rninutus genibarbis group: Skutch 1969, V80, D. Robinson; X. rn. minutus: Euler 1900; X. rutilans: Belcher and Smooker 1936 (HMNH 11677-8), Willis 1988, KZ.

APPENDIX2. Distributionof statesof 24 nest-architecturecharacters in 168 speciesof furnariidsand two compositeoutgroups. Furnariid taxa are condensed into 49 OTUsand listed in alphabeticalorder. Numbers in parenthesesrefer to thenumber of specieswithin a givenOTU. 0 = ancestralstate; 1, 2 = derivedstates; P = polymorphism(i.e. 0 or 1); ? = unknown.See text for characterdescriptions and polarity assessments and Appendix1 for sourcesof nestinformation.

Characters

Taxa a.b 1-5 6-10 11-15 16-20 21-24 Acrobatornisfonsecai 00000 00000 10011 00100 0000 Anabazenops(2) 12000 01000 00000 00000 0000 Anumbius annumbi 00000 00000 10011 01000 0000 Aphrasturaspinicauda 10000 00000 10000 00000 0000 Asthenes anthoides 00000 00000 10011 00000 0000 Astheneshumicola group (11) 00000 00000 10011 00100 0000 Asthenespyrrholeuca group (2) P0000 00000 100 P1 00 P00 0000 Asthenes wyatti group (7) 00000 00000 3.0000 00000 0000 Autornolusleucophthalrnus group (3) 3.3. 000 3.0000 00000 00000 0000 Automolus rubiginosus 12000 02000 00000 00000 0000 Chilia rnelanura 3_3_001 00000 00000 00000 0000 Certhiaxis (2) 00000 00000 1003_2 3_03_00 0000 Cinclodes (11) 12000 01000 00000 00000 0000 Coryphistera alaudina 00000 00000 10011 01000 0000 Cranioleuca albiceps group (11) 00000 00000 10000 00000 0010 Cranioleuca pyrrhophia group (7) 00000 00000 10000 00000 0000 Eremobius phoenicurus 00000 00000 10011 00100 0000 Furnarius (6) P2000 01001 00000 00000 0000 October1999] NestPhylogeny ofFurnariidae 911

APPENDIX 2. Continued.

Characters

Taxa a,b 1-5 6-10 11-15 16-20 21-24

Geositta (9) 12000 01000 00000 00000 0000 Gyalophylax hellmayri 00000 00000 10012 10100 0000 Hylocryptus (2) 12000 02000 00000 00000 0000 Hyloctistes subulatus 11000 10000 00000 00000 0000 Leptasthenura (9) 70000 00000 lOOO0 00000 0000 Limnoctites rectirostris 00000 00000 10000 00000 1000 Limnornis curvirostris 00000 00000 10000 00000 1000 Lochmias nematura 10000 00000 10000 00000 0000 Margarornis (2) 00000 00000 10000 00000 0010 Ochetorhynchus certhioides 12000 01100 00000 00000 0000 Oreophylax moreirae 00000 00000 10100 00000 0000 Phacellodornus (8) 00000 00000 10013_ 00101 0001 Philydor (4) 12000 02000 00000 00000 0000 Phleocryptesmelanops 00000 00000 11000 00000 1100 Poecilurus (2) 00000 00000 10012 10100 0000 Premnoplex brunnescens 70000 00000 10000 00000 0010 Pseudocolaptes (2) 11010 00000 00000 00000 0000 Pseudoseisura (3) 00000 00000 10012 07100 0000 Pygarrhichas albogularis 11100 00010 00000 00000 0000 Schizoeaca (4) 00000 00000 10100 00000 0000 Schoeniophylaxphryganophila 00000 00000 10012 10100 0000 Sclerurus (6) 11000 10000 00000 00000 0000 Siptornis striaticollis 00000 00000 10000 00000 0010 Spartonoica maluroides 00000 00000 10000 00000 0000 Sylviorthorhynchusdesmursii 00000 00000 10000 00000 0000 Synallaxis (19) 00000 00000 20012 10100 0000 Thripadectes melanorhynchus 11000 10000 00000 00000 0000 Thripadectesrufobrunneus group (2) 12000 02000 00000 00000 0000 Thripophagafusciceps 00000 00000 10000 00010 0010 Upucerthia (7) 12000 01000 00000 00000 0000 Xenops (3) 12000 02010 00000 00000 0000 Dendrocolaptidae l1000 00000 00000 00000 0000 Formicarioidea PP000 PP000 P0000 00000 0000

aSpecies representing composite OTUs (asterisksmark specieswith partial information):Anabazenops = A. dorsalisand A. fuscus*;Asthenes hurnicola group = A. arequipae, A. baeri, A. berlepschi*, A. cactorutn,A. dorbignyi, A. huancavelicae,A. hutnicola,A. luizae*, A. patagonica. A. pudibunda, and A. steinbachi;Asthenes pyrrholeuca group = A. tnodestaand A. pyrrholeuca; Astheneswyatti group = A. hudsoni,A. hutnilis, A. tnaculicauda,A. sclateri, A. virgata, A. urubatnbensis*,and A. wyatti; Automolusleucophthalmus group = A. infuscatus,A. leucophthaltnus, and A. ochrolaetnus; Certhiaxis = C. cinnamotnea and C. mustelina; ( inclodes = C. antarcticus, C. atacatnensis, C. comechingonus,C. excelsior, C. fuscus, C. nigrofutnosus,C. olrogi, C. oustaleti,C. pabsti, C. patagonicus,and C. taczanowskii*;Cranioleuca albiceps group = C. albicapilla, C albiceps, C. antisiensis,C. baroni, C. curtara, C. detnissa,C. erythrops,C. helltnayri, C. tnarcapatae.C. subcristata,and C. vulpina; Cranioleuca pyrrhophia group = C. gutturata, C. tnuelleri, C. obsoleta, C. pallida, C. pyrrhophia, C. setnicinerea,and C. sulphurifera;Furnarius = F. cristatus, F. figulus, F. leucopus,F. tninor, F. rufus, and F. torridus; Geositta = G. antarctica, G. cunicularia, G. isabellina*. G. tnarititna*, G. peruviana, G. punensis*, G. rufipennis,G. saxicolina*. and G. tenuirostris;Itylocryptus = H. erythrocephalusand H. rectirostris;Leptasthenura = L. aegithaloides,L. andicola, L. fuliginiceps, L. pileata, L. platensis,L. sertaria, L. striolata, L. xenothorax,and L. yanacensis;Margarornis • M. rubiginosusand M. squatniger;Phacellodomus = P. dorsalis*, P. erythrophthaltnus,P. tnaculipectus,P. ruber, p. rufifrons, P. sibilatrix, P. striaticeps,and P. striaticollis; Philydor = P. atricapillus, P. erythrocercus*,P. lichtensteini*,and P. rufus; Poecilurus = P. candei and P. scutatus; Pseudoseisura = P. cristata, P. gutturalis, and P. lopbores; Pseudocolaptes • P. boissonneautii* and P. lawrencii; Schizoeaca = S. fuliginosa, S. griseotnurina*, S. harterti, and S. helleri; Sclerurus = S. albigularis, S. caudacutus,S. guatetnalensis,S. tnexicanus,S. rufigularis. and & scansor;Synallaxis = S. albescens,S. albigularis, S. azarae, & brachyura, S. chinchipensis,S. cinerascens,S. cinnamomea,S. erythrothorax, S. frontalis, S. gujanensis,S. tnacconnelli*,S. propinqua*, S. ruficapilla, S. rutilans, S. spixi, S. stictothorax,S. subpudica*,S. tithys, and S. zitntneri; Thripadectes rufobrunneus group = T. rufobrunneusand T. virgaticeps; Upucerthia = U. albigula. U. andaecola, U. dutnetaria, U. jelskii*, U. ruficauda*, U. serrana*, and U. validirostris; Xenops = X. tnilleri*, X. minutus, and X. rutilans. bThe genera Anabacerthia, Berlepschia, Geobates, Metopothrix, Siptornopsis,Syndactyla. and Xenerpesteswere not included in the analysis becausenest data were insufficient for characterstates to be coded accurately.