PHYLOGENY OF THE TYRANT FLYCATCHERS (TYRANNIDAE) BASED ON

715

The Auk 119(3):715–734, 2002

PHYLOGENY OF THE TYRANT FLYCATCHERS (TYRANNIDAE)BASED ON MORPHOLOGY AND BEHAVIOR

JEFFREY S. BIRDSLEY1

W. K. Kellogg Biological Station and Department of Zoology, Michigan State University, 3700 East Gull Lake Drive,Hickory Corners, Michigan 49060, USA

ABSTRACT.—Previously published morphological and behavioral data for the tyrant fly-catchers (Tyrannidae) were reanalyzed using cladistic techniques. Several additional char-acters, including two putative synapomorphies of the Tyrannidae were incorporated. Nearlyall of the ;100 traditional tyrannid genera were included. Results of the analysis of this setof 68 characters support three previously proposed tyrannid assemblages: the kingbird as-semblage and slightly restricted Empidonax and Myiarchus assemblages. Characters of thenasal septum that past workers have considered conservative and phylogenetically infor-mative are supported as synapomorphies of the kingbird and restricted Empidonax assem-blages. Several small monophyletic groups of genera are supported. The monophyly of nei-ther the Elaenia assemblage nor the flatbill and tody-tyrant assemblage is supported in anymost-parsimonious tree, but such trees are not significantly better than trees supportingtheir monophyly. A Tyrannidae exclusive of the Cotingidae and Pipridae is monophyletic insome most-parsimonious trees, but not in others in which flatbills and tody-tyrants are basalto a clade containing Cotingidae, Pipridae, and the remaining tyrannid genera. There is someevidence that characters describing plumage color pattern are more homoplasious than theother character suites I examined. Received 5 August 2000, accepted 9 February 2002.

RESUMEN.—Utilizando tecnicas cladısticas reanalice datos morfologicos y conductualespreviamente publicados para la familia Tyrannidae. Incorpore varios caracteres adicionalesincluyendo dos sinapomorfias putativas. Incluı casi todos los generos de los aproximada-mente 100 generos tradicionales de la familia Tyrannidae. Los resultados del analisis de unset de 68 caracteres apoyan tres grupos de tiranidos previamente propuestos: el grupo Tyran-nus y los grupos Emipidonax y Myiarchus. Los caracteres relacionados con el septum nasalque han sido considerados previamente como conservadores y filogeneticamente informa-tivos, son apoyados como sinapomorfias del grupo Tyrannus y del grupo restringido Empi-donax. Varios pequenos grupos de generos monofileticos son apoyados. Ni la monofilia delgrupo Elaenia ni la de los picochato (e.g. Plathyrhinchus) y mosqueritos (e.g. Todirostrum , He-mitriccus) es apoyada en cualquiera de los arboles mas parsimoniosos, pero estos arboles noson significativamente mejores que los que apoyan su monofilia. Un Tyrannidae exclusivode la familia Cotingidae y Pipridae es monofiletico en alguno de los arboles mas parsimo-niosos, pero no en otros en los que los picochatos y mosqueritos son basales a un clado quecontiene a Cotingidae, Pipridae y a los demas generos de tiranidos. Existe cierta evidenciaque los carateres que describen los patrones de coloracion del plumaje son mas homoplasicosque el set de caracteres que examinee.

THE TYRANT FLYCATCHERS (Tyrannidae) are alarge, primarily Neotropical family of subos-cine passerine birds in the superfamily Tyran-noidea which also include the cotingas (Cotin-gidae) and manakins (Pipridae). A greatbehavioral and ecological diversity within thatsuperfamily presents rich opportunities forcomparative tests of evolutionary and ecologi-

1 Present address: Department of Biological Sci-ence, Florida State University, Tallahassee, Florida32306, USA. E-mail: [email protected]

cal hypotheses (e.g. Prum 1990a, 1994, 1997).Because well-supported phylogenies are fun-damental to such tests, the evolutionary rela-tionships of these birds have received much re-cent attention (W. E. Lanyon 1984, 1985, 1986,1988a, b, c; McKitrick 1985; S. M. Lanyon 1985;Prum and Lanyon 1989; Prum 1990b, 1992; Mo-bley and Prum 1995; Bostwick 2000). W. E. Lan-yon’s work in particular has resulted in an ex-tensive comparative morphological andbehavioral data set for the Tyrannidae.

The relationships among the 90 to 100 generaof tyrant flycatchers were historically based on

716 [Auk, Vol. 119JEFFREY S. BIRDSLEY

the shape of the bill, wing and tail, and colorand pattern of plumage. Warter’s (1965) surveyof cranial osteological variation in the Tyran-noidea revealed variation in the nasal capsuleand septum, palatine and palatomaxillarybones, interorbital septum, and general shapeof the cranium. Warter felt that the nasal regionprovided useful taxonomic characters and rec-ognized several basic states of the configura-tion of the nasal septum. Building upon War-ter’s (1965) work, W. E. Lanyon (1984, 1985,1986, 1988a, b, c) completed a nearly exhaus-tive survey of tyrannoid cranial variation andused states of the nasal septum to hypothesizemonophyly of five separate tyrannid assem-blages that collectively included nearly all tra-ditional tyrannid genera.

Ames (1971) documented great variation insuboscine syringeal musculature and supportelements. However, Ames did not use differ-ential stains and so could not readily distin-guish between ossified and cartilaginous sup-port elements. Lanyon (1984, 1985, 1986, 1988a,b, c) studied a nearly exhaustive sample oftyrannoid syringes which he stained to distin-guish the ossified (primarily tracheal) series ofelements from the cartilaginous (primarilybronchial) series. This allowed recognition ofhomologous elements. Lanyon used syringealcharacters to group taxa within his five assem-blages and used characters of the cranium,nesting behavior, plumage, and egg colorationwherever necessary to establish relationshipswithin syringeal groups and occasionally athigher levels. That procedure is the equivalentof weighting those syringeal characters moreheavily than others, effectively disallowingconvergence in them. Nasal septum characterswere effectively weighted even more heavilythan syringeal characters because Lanyon as-sumed them to be synapomorphies of his fiveassemblages and disallowed convergence inthem.

Lanyon (1984, 1985, 1986, 1988a, b, c) devel-oped hypotheses of generic relationships basedon shared similarities, but because he did notdo so within a cladistic framework, some of hisgroups were supported by plesiomorphies. Forexample, in the Elaenia group Lanyon (1988a)used the poorly ossified condition of the ante-rior segment of the nasal septum to support aclade composed of Capsiempis, Phaeomyias, andNesotriccus. He then used the fully ossified con-

dition to support a clade composed of Serpop-haga and Anairetes. Only one of those characterstates is the derived state and the other is theplesiomorphic, or ancestral, state in that group.Logically, only the derived state may argue forclose relationship of the taxa that possess it.

A cladistic reanalysis of Lanyon’s (1984,1985, 1986, 1988a, b, c) data was performedplus any other data that could be found in theliterature. I weighted characters equally andanalyzed them together in one character matrixcontaining nearly all tyrannid genera, includ-ing several that Lanyon could not place in anyof his assemblages. That provided a test of themonophyly of each of the five tyrannid assem-blages Lanyon proposed and provided a test ofhomology of each of the five nasal septumstates that supported them. Monophyly of theTyrannidae has been questioned on the basis ofDNA–DNA hybridization analyses that placeseveral ‘‘mionectid’’ flycatchers as the sistergroup to the rest of the tyrannoids (cotingids 1piprids 1 remaining tyrannids) (Sibley andAhlquist 1985). The present analysis provides atest of the monophyly of the Tyrannidae as tra-ditionally constituted (e.g. American Ornithol-ogists’ Union [AOU] 1998).

I compared reliability of morphological, be-havioral, and ecological characters to recovertyrannid phylogeny. Although past workershave argued (e.g. Atz 1970) that behavioral andecological characters lack phylogenetic infor-mation, several studies have found such char-acters to be at least as reliable as morphologicaland molecular characters (McLennan and Mat-tern 2001, Paterson et al. 1995, DeQueiroz andWimberger 1993, but see Lee et al. 1996). Sig-nificantly different levels of homoplasy amongfour suites of tyrannid characters was testedfor: cranial, syringeal, plumage, and behavioraland ecological characters.

METHODS

All 95 genera that served as terminal taxa in Lan-yon’s (1984, 1985, 1986, 1988a, b, c) five assemblageswere included and the nomenclature follows his.Lanyon provided character support for monophylyof 64 of those genera. Lanyon (1986, 1988a) split Me-cocerculus and Myiophobus into three groups each andplaced them in different parts of his phylogenies yetretaining their generic names until more compara-tive data are available. Here these taxa are split asLanyon did. Lanyon (1988b) argued for merging ’’T-erenotriccus’’ erythrurus into Myiobius and his rec-

July 2002] 717Phylogeny of Tyrannidae

FIG. 1. Character state tree for character 11. Char-acters and states are described in Appendix 1.

ommendation are followed here, but Myiobius eryth-rurus is kept as a terminal taxon separate from theremaining Myiobius. Several traditional tyrannidtaxa have a nasal septum morphology that is eitherunknown or is sufficiently unique that Lanyon (1986,1988a) refrained from placing them in any of his as-semblages, maintaining them incertae sedis: Phyllom-yias fasciatus, P. griseiceps, P. griseocapilla, and themonotypic genera Colonia, Culicivora, Machetornis,Muscigralla, and Tachuris. All of these taxa are in-cluded in this analysis except for Phyllomyias griseo-capilla and Culicivora. For those two taxa, neither cra-nial nor syringeal data were available. Also includedis the genus Neopipo, traditionally placed in the Pi-pridae, because Mobley and Prum (1995) hypothe-sized it to belong in the Tyrannidae on the basis ofsyringeal and plumage data.

Character data were extracted primarily from Lan-yon (1984, 1985, 1986, 1988a, b, c). These referencesinclude photographs of representative syringes forall terminal taxa and photographs of crania for most.Because Lanyon reported on one assemblage at atime, he often did not explicitly describe, for a givencharacter, the distribution of states across all tyran-nid genera and in outgroups. In those cases, family-wide assessments and outgroup comparisons weremade using Lanyon’s photographs and other litera-ture sources describing tyrannoid crania (Warter1965) and syringes (Ames 1971; McKitrick 1985;Prum and Lanyon 1989; Prum 1990b, 1992; Mobleyand Prum 1995). Lanyon did not make extensive useof syringeal musculature characters so those wereextracted from Ames (1971). Because Ames (1971)did not use stains on ossified and cartilaginous sy-ringeal support elements precedence was given to allmore-recent information on those elements. Hiltyand Brown (1986), Stiles and Skutch (1989), Ridgelyand Tudor (1994) and similar guides for plumagecharacters were consulted. Foraging behavior char-acters are from Fitzpatrick (1980). For nest and eggcharacters, an exhaustive literature search was con-ducted, drawing information from many sources andadding two characters that Lanyon did not use: pres-ence of a visor over the entrance of enclosed nests(character 66) and nests used as a dormitory (char-acter 67). In addition, crania, skins, and Lanyon’scleared-and-stained syringeal specimens were ex-amined at the American Museum of Natural Historyfor any character data that were incomplete or in-conclusive in the literature.

Lanyon (1984, 1986, 1988a) hypothesized theTyrannidae to be monophyletic based on the pres-ence of internal syringeal cartilages. Prum (1990b)questioned the homology of all internal syringealcartilages and recommended using unique, detailedmorphologies of internal cartilages as characters,while refraining from the hypothesis of homology ofinternal cartilages as broadly defined. Prum (1990b)hypothesized monophyly of the Tyrannidae on the

basis of the presence of Mm. obliqui ventrales, a pairof intrinsic syringeal muscles. In this analysis ’’Mm.obliqui ventrales’’ is included as a character but not‘‘internal cartilages,’’ except as several charactersthat describe unique details of some internal carti-lages. An additional, heretofore unrecognized, po-tential synapomorphy of the tyrannids is included:B1 and B2 syringeal support elements which are con-nected at their ventral tips. That connection is absentin cotingids and piprids (Prum 1992; R. O. Prumpers. comm.).

Morphological and behavioral variation were cod-ed as 50 binary characters, 15 unordered multistatecharacters, and 3 ordered multistate characters (Ap-pendix 1). Characters 6 and 13 were ordered becausestates shared significant detail but some states hadadditional details that appeared to be further de-rived. For character 11 a state tree (Fig. 1) is definedreflecting a trend toward increasing numbers of dou-ble, complete syringeal A elements and increasingdegree of ossification of those elements. That wasjustified because such a trend has apparently oc-curred in piprids (Prum 1992) and in some falconi-forms (Griffiths 1994). Also, the ontogenetic devel-opment of those elements proceeds from an entirelycartilaginous state to an at least partly ossified state(Ames 1971).

PAUP* 4.0b8 (Swofford 2000) was used to performheuristic searches for the most-parsimonious trees.To increase the probability of finding globally most-parsimonious trees, I repeated this search using allpossible combinations of branch-swapping and step-wise-addition options. For each search I set a maxi-mum of 2,000 trees to be held in memory. I also per-formed 1,000 replicates with the random additionoption and one tree held in memory. Multistate taxawere interpreted as polymorphisms. I gave all char-acters equal weight. I felt this was justified in the ab-sence of objective criteria for weighting some char-acters more heavily than others. Equal weightingschemes do, however, carry the assumption of lowrates of change in all characters (Felsenstein 1982). Irooted all trees to Furnarioidea and allowed the oth-er potential outgroups, Cotingidae and Pipridae, to‘‘float’’ in the analysis. This allowed a test of themonophyly of the Tyrannidae as traditionallyconstituted.


I used consensus trees to summarize the resultingset of most-parsimonious trees. I computed a strictconsensus tree which identifies groups found in allmost-parsimonious trees. I also computed an Adamsconsensus tree (Adams 1972) which better illustratespatterns and trends among alternative most-parsi-monious trees by identifying groups (not necessarilymonophyletic) that consistently nest within largergroups in all trees. For example, if a set of nestedgroups exists in all most-parsimonious trees but insome of them is ‘‘invaded’’ by one taxon or a smallgroup of taxa, the Adams consensus preserves thenested structure and places the problematic groupoutside of it in a polytomous position. The strict con-sensus simply collapses the entire nested group.Consensus trees should be interpreted with care asthey may not be the most parsimonious hypothesessuggested by the data (Swofford 1991).

To evaluate the evidential support for relation-ships depicted in the strict consensus tree I used So-renson’s (1996) TreeRot program and PAUP* to com-pute a decay index (Bremer 1988, Kallersjo et al.1992). I ran 10 replicates with TBR branch-swapping,random addition sequence and a maximum of 800trees held in memory. I also ran 100 replicates withrandom addition sequence and one tree held inmemory. The decay index gives the number of extrasteps required to show a node as unsupported. Forexample, if a node that occurs in all most-parsimo-nious trees is not present in trees that are one steplonger, that node gets a decay index of 1. Higher in-dex values indicate more robust clades.

I used two approaches to evaluate Lanyon’s (1984,1985, 1986, 1988a, b, c) hypothesized assemblageswithin the context of my data matrix. First, for thoseassemblages not supported in the strict consensus ofmost-parsimonious trees, I determined the numberof extra steps required to force their monophyly. Todo this I conducted heuristic searches constrainingmonophyly of each assemblage in turn. For each I ran10 replicates with TBR branch-swapping, randomaddition, and a maximum of 800 trees held in mem-ory. I also ran 1,000 replicates with one tree held inmemory. I used Templeton’s test (Templeton 1983) asimplemented in PAUP* to determine whether the re-sulting trees were significantly different from the most-parsimonious trees. Second, I used MACCLADE 3.04(Maddison and Maddison 1992) to examine evolu-tion of nasal septum characters on the most-parsi-monious trees. I evaluated the homology of septumcharacters that Lanyon (1984, 1985, 1986, 1988a, b, c)used to define his assemblages.

Finally, I used MACCLADE to calculate rescaledconsistency indices (RC) for each character and to av-erage them across 2,000 most-parsimonious treesrepresenting all known tree islands. I used a singlefactor ANOVA to determine if mean RC was signifi-cantly different among four suites of characters: cra-nial (characters 1–10), syringeal (characters 12–45),

plumage (characters 46–59), and behavior/ecology(characters 60–67). I did not include character 11 be-cause MacClade does not include characters of user-defined type in these calculations.

RESULTS AND DISCUSSION

Phylogenetic analysis of 68 characters (a totalof 101 apomorphies) and 106 taxa (Appendix 2)identified more than 2,000 most-parsimonioustrees of length 514 and a consistency index (CI)of 0.44. I found several ‘‘islands’’ of most-par-simonious trees using different combinationsof PAUP’s search options. I found additionaltree islands in the course of computing decayindices. The strict consensus of all most-parsi-monious trees (Figs. 2–4) indicates support forthe monophyly of many small groups of generaand of three assemblages which correspondmore or less to Lanyon’s (1984, 1985, 1986)kingbird, Myiarchus, and Empidonax assemblag-es. The monophyly of neither an Elaenia assem-blage (Lanyon 1988a) nor a flatbill and tody-ty-rant assemblage (Lanyon 1988c) is supported.An Adams consensus tree (Figs. 5 and 6) is bet-ter resolved. Each clade depicted in the Adamstree is present in at least some most-parsimo-nious trees, but the tree as a whole may not beconsistent with any single most-parsimonioustree. Despite this, for the purposes of the fol-lowing discussion it illustrates many resultswhich are not apparent in the strict consensus(Figs. 2–4). Here I discuss my results as com-pared to each of Lanyon’s (1984, 1985, 1986,1988a, b, c) proposed assemblages and as com-pared to the traditional concept of the familyTyrannidae. Character numbers refer to char-acters in the present analysis.

Tyrannidae monophyly. A traditional Tyran-nidae exclusive of Cotingidae and Pipridae issupported in some most-parsimonious trees.However, there are no unambiguous synapo-morphies. Neither of the two putative synapo-morphies of the Tyrannidae provides unequiv-ocal support for monophyly of the family astraditionally defined. In all most-parsimonioustrees, intrinsic syringeal muscles Mm. obliquiventrales that insert ventrally (character 43,state 1) are a synapomorphy of a restrictedTyrannidae whose exact membership variesfrom tree to tree but never includes the tody-tyrant group or the Cnipodectes–Onychorhyn-chus–Myiobius clade. The ventrally connectedB1 and B2 elements (character 19) proved dif-


FIG. 2. Strict consensus tree derived from allmost-parsimonious trees. Decay indices are shownwhen greater than 1. The two taxa at the bottom de-note the assemblages shown in Figures 3 and 4.

FIG. 3. Strict consensus tree of the kingbird as-semblage. Decay indices are shown when greaterthan 1.

ficult to evaluate as a synapomorphy of theTyrannidae. The basal or next-to-basal groupsin each most-parsimonious tree possess thischaracter, but the outgroup (Furnarioidea) wascoded ’’?’’ (unknown). Therefore character 19does not provide an unequivocal synapo-morphy of the Tyrannidae in any most-parsi-monious trees. To assess the level at which it isinformative, one would need more informationon this trait in the Furnarioidea and Old Worldsuboscines.

Flatbill and tody-tyrant assemblage. Lanyon(1988c) defined a flatbill and tody-tyrant as-semblage based on the possession of: unossi-fied nasal septa (character 1); cranial interor-bital septa in which the supraorbital fenestra is

reduced (character 9); and enclosed, pendantnests (character 64). This assemblage is notsupported in any most-parsimonious trees.Four extra steps are required to force its mono-phyly, but such trees are not significantly dif-ferent from most-parsimonious trees (P . 0.64;21 , n , 27 characters requiring differingnumbers of steps on the trees being compared).Within his assemblage, Lanyon (1988c) defineda flatbill group (Rhynchocyclus, Tolmomyias,Onychorhynchus, and Platyrinchus) based on thepossession of at least one ossified, double, com-plete syringeal A element (character 11). Thisstate is present in many tyrannids outside theflatbills and tody-tyrants and is not a synapo-morphy of these flatbills in any most-parsi-monious trees. Lanyon’s (1988c) hypothesizedsister relationship between Onychorhynchus andPlatyrinchus is not supported in any most-par-simonious trees. Two seemingly unique syrin-geal characters (23 and 31) shared by Ony-


FIG. 4. Strict consensus tree of the restricted Em-pidonax assemblage. Decay indices are shown whengreater than 1.

FIG. 5. Adams consensus tree derived from allmost-parsimonious trees. The taxon at the bottomdenotes the assemblage shown in Figure 6.

chorhynchus and Platyrinchus may be derivedindependently in these two taxa. A sister rela-tionship between Rhynchocyclus and Tolmomyiasis supported in all most-parsimonious trees bythe presence of narrow, linear internal syrin-geal cartilages (character 29), a retort-shapedpendant nest (character 65) which is used as adormitory (character 67). In all most-parsimo-nious trees this relationship is additionallysupported by at least two of the following char-acters: 1, 11, 43, 49, 64.

Lanyon (1988c) could not determine the re-lationships of Cnipodectes within the assem-blage, maintaining it in a polytomy with hisflatbill clade and tody-tyrant clade. My resultssuggest that Cnipodectes is the sister group to aclade composed of Onychorhynchus and Myio-bius (including erythrurus). This relationship issupported in all most-parsimonious trees by

the presence of long rictal bristles (character 57)and rufous tails (character 53; lost within Myio-bius). I discuss the hypothesized sister relation-ship between Onychorhynchus and Myiobius be-low with the Empidonax assemblage.

Lanyon (1988c) defined a tody-tyrant group(Todirostrum, Poecilotriccus [including Taenio-triccus], Hemitriccus [including Myiornis], Lop-hotriccus [including Atalotriccus], and Oncosto-ma) based on the possession of two characters:1) a horseshoe-shaped, cartilaginous bronchialplate, and 2) delicate, rod-like internal cartilag-es located near the caudal ends of the horse-shoe. In the present study, I combined theseinto one character state (character 27, state 1)which supports monophyly of Lanyon’s tody-tyrant group in some, but not all most-parsi-monious trees. Lanyon (1988c) proposed aclade composed of Hemitriccus, Lophotriccus,and Oncostoma based on the presence of a pes-sulus that extends anteriorly to divide four ormore A elements dorsally (character 16). Thisclade is supported in all most-parsimonioustrees by character 16 and, additionally in somemost-parsimonious trees, by characters 43 and


FIG. 6. Adams consensus tree of an ‘‘elaeniid andempid’’ clade.

64. Lophotriccus and Oncostoma are supportedas sister groups by a laterally compressed syr-inx and trachea (character 26) in all most-par-simonious trees.

There is some support for a basal placementof the flatbills and tody-tyrants within theTyrannidae and perhaps within the Tyrannoi-dea. In all most-parsimonious trees the flatbillsand tody-tyrants are basal to a large clade con-taining the majority of tyrannids plus, in somemost-parsimonious trees, cotingids and pi-prids. This large clade is supported by thepresence of ossified nasal septa (character 1).This is at least partly congruent with the resultsof Sibley and Ahlquist’s (1985) DNA–DNA hy-bridization analyses in that the tody-tyrants To-dirostrum and Hemitriccus were among the taxathey placed in their family ‘‘Mionectidae’’ asthe sister group to the rest of the tyrannoids(cotingids 1 piprids 1 remaining tyrannids).

Kingbird assemblage. Results support akingbird assemblage completely consistent

with that of Lanyon (1984) (Fig. 3). It is sup-ported in all most-parsimonious trees by char-acters 3, 9, 10, 44, and 55. My hypothesis of re-lationships within this assemblage is, however,less well resolved than Lanyon’s (1984). This isdue to the fact that Lanyon (1984) used alter-native derived states of the ventral connectionof the B1 and B2 syringeal support elements(character 19) to support two major cladeswithin the assemblage: Phelpsia, Pitangus, Phi-lohydor, Legatus, and Myiozetetes have flattenedconnections (state 1) whereas the remainingseven genera in the assemblage have roundedconnections (state 2). My family-wide analysisindicates that flattened connections, which oc-cur in nearly all tyrannids, are ancestral in thekingbird assemblage and thus do not argue forrelationships within it. My results do not sup-port monophyly of Lanyon’s clade composed ofthe five genera named above. However, round-ed connections (state 2) appear to be unique toMyiodynastes, Conopias, Megarynchus, Tyrannop-sis, Tyrannus, Empidonomus, and Griseotyrannus.This clade is supported in all most-parsimo-nious trees by characters 19 and 32. Lanyon’shypothesized relationships among these gen-era are also supported in all most-parsimoni-ous trees. Character 62 supports a sister-grouprelationship between Myiodynastes and Cono-pias. Characters 20, 42, and 61 support a cladecomposed of Megarynchus, Tyrannopsis, Tyran-nus, Empidonomus, and Griseotyrannus. Charac-ter 32 supports a sister-group relationship be-tween Megarynchus and Tyrannopsis. Character59 supports a clade composed of Tyrannus, Em-pidonomus, and Griseotyrannus.

Character 11 supports sister-group relation-ships between Pitangus and Philohydor (two ormore double, complete, ossified A elements)and between Legatus and Myiozetetes (incom-plete A3 elements fused to complete A2). Char-acter 45 (intrinsic syringeal muscles Mm. obli-qui laterales) supports the Legatus–Myiozetetesrelationship in some most-parsimonious trees,but in the remainder of trees, this is ambiguousdue to uncertainty about the state of this char-acter in Phelpsia.

Empidonax assemblage. A restricted Empido-nax assemblage (Fig. 4) is supported contain-ing the same genera that Lanyon (1986, 1988b)placed in it except for the four genera that com-prise his Myiophobus group: Myiophobus,Pyrrhomyias, Hirundinea, and Myiobius. Seven


extra steps are required to force a monophyleticEmpidonax assemblage including the Myiopho-bus group. However, such trees are not signifi-cantly different from most-parsimonious trees(P . 0.36; 14 , N , 26 characters requiring dif-fering numbers of steps on the trees being com-pared). In all most-parsimonious trees themonophyly of the restricted Empidonax assem-blage is supported by the presence of a nasalseptum with basal trabecular plate and ante-rior notch (character 4), a wide posterior forkon the trabecular plate (character 7), and ‘‘near-ground generalist’’ foraging behavior (charac-ter 61). In all most-parsimonious trees, the‘‘Myiophobus group’’ genera are distantly relat-ed to this restricted Empidonax assemblage.

Within the Empidonax assemblage, these re-sults support Lanyon’s (1986) hypothesis of abasal position of the chat-tyrants Ochthoeca, Sil-vicultrix and Colorhamphus. These results do notsupport the monophyly of Lanyon’s (1986)Ochthoeca group composed of these three gen-era plus Arundinicola, Fluvicola, and Alectrurus.Lanyon (1986) used character 11, state 0 (no Aelements completely encircle the bronchi) tosupport his Ochthoeca group, but my family-wide analysis indicates that this state is ances-tral in the Empidonax assemblage and does notargue for a close relationship of these taxa.

The monophyly of a large clade (Alectrurusthrough Lathrotriccus in Fig. 4) composed of allof the restricted Empidonax assemblage generaexcept Ochthoeca, Silvicultrix and Colorhamphusis supported in all most-parsimonious trees bycharacters 22, 49, or both. Within this largeclade, a group (Cnemotriccus to Lathrotriccus inFig. 4) similar to Lanyon’s (1986) Empidonaxgroup is supported in all most-parsimonioustrees by character 49, and additionally in someof those trees by characters 13, 22, and 61. Sa-yornis, which possesses the uniquely modifiedsyringeal A2 elements (character 13) of thisgroup is, in some most-parsimonious trees, thebasal member of the group. In other most-par-simonious trees, however, Sayornis groups withother taxa that lack wing bars (character 49)and are ‘‘near-ground generalist’’ foragers(character 61). In all most-parsimonious trees,characters 24, 25, and 52 support Gubernetesand Muscipipra as sister groups.

Few additional relationships are well re-solved in the strict consensus of this assem-blage (Fig. 4), but upon examination of the set

of most-parsimonious trees and of the Adamsconsensus (Fig. 6), some patterns are evident.For example, all most-parsimonious trees con-tain a clade corresponding roughly to Lanyon’s(1986) Muscisaxicola group containing theground-tyrants, shrike-tyrants, bush-tyrantsand allies. Hymenops, which shares some plum-age and habitat characters with this group, is,in some most-parsimonious trees, placed with-in this clade (Muscisaxicola through Cnemarchusin Fig. 6); hence, a Muscisaxicola group is notdepicted in the strict consensus (Fig. 4). Thisgroup, with or without Hymenops, is supportedby characters 60 (habitation of open grasslandor marsh) and 61 (ground specialist foragingbehavior). In agreement with Lanyon’s (1986)concept of this group, Muscisaxicola always isplaced basally, as illustrated in Figure 6. Pat-terned wings (character 48) and notched or at-tenuated primaries (character 59) supportmonophyly of the rest of the group exclusive ofMuscisaxicola. ‘‘Perch-to-ground’’ foraging be-havior (character 61) supports a group exclu-sive of Muscisaxicola and Lessonia.

A clade composed of Agriornis, Myiotheretes,Polioxolmis and Cnemarchus is supported in allmost-parsimonious trees by the presence ofstreaked throat plumage (character 50). Lan-yon’s (1986) hypothesized bush-tyrant cladecomposed of Myiotheretes, Polioxolmis and Cne-marchus is supported in all most-parsimonioustrees by an ossified interorbital septum (char-acter 9) and a transition from open grasslandto forest–woodland habitat (character 60). Lan-yon (1986) was unable to resolve relationshipsamong these genera, but these results supportPolioxolmis and Cnemarchus as sister groupsbased on the relatively narrow dorsal ends oftheir B1 syringeal support elements (character22, state 0), and the presence of well developedMm. obliqui ventrales (character 44). Lanyon(1988a) used character 44 in the Elaenia assem-blage but apparently did not recognize it in theEmpidonax assemblage. Lanyon (1986) hypoth-esized a clade composed of Arundinicola, Flu-vicola, Alectrurus supported by characters 8, 56,and 60. In the present study, these characterssupport such a clade in some most-parsimoni-ous trees. The remainder of most-parsimonioustrees hypothesize an (Arundinicola (Fluvicola(Alectrurus, Hymenops))) clade. Hymenopsshares plumage and habitat characters with theother three genera, but Lanyon (1986) placed


Hymenops in his Knipolegus group with which italso shares plumage and syringeal characters.Arundinicola, Fluvicola, and Alectrurus, at least,appear to be closely related, as depicted in Fig-ure 6.

The double, complete, ossified syringeal A el-ements (character 11) and concealed crownpatch (character 46) present in Lanyon’s (1986,1988b) Myiophobus group are not present in therest of the Empidonax assemblage and argue forthe placement of these genera elsewhere. In theset of most-parsimonious trees, Myiophobus,Pyrrhomyias, and Hirundinea are allied withseveral different groups outside the Empidonaxassemblage. A sister-group relationship be-tween Myiobius (including erythrurus) and Ony-chorhynchus is supported by the presence of atleast two double, complete, ossified syringeal Aelements (character 11) and an ossified inter-orbital septum (character 9). Myiobius also hasthe long rictal bristles (character 57) and eryth-rurus has the rufous tail (character 53) found inOnychorhynchus and Cnipodectes. Past workers(Ames 1971, Traylor 1977, Traylor and Fitzpat-rick 1982) have hypothesized a close associa-tion between Myiobius and Onychorhynchusbased on these syringeal and plumage charac-ters. Myiobius builds a pendant, enclosed nest(character 64) as do Onychorhynchus and Cni-podectes, but in the present analysis this char-acter does not support the relationship of Myio-bius to these genera because the globular nests(state 1) of Myiobius and elongate nests (state 2)of Onychorhynchus and Cnipodectes were codedas alternative unordered states. Characters 1, 4,19, and, in some most-parsimonious trees, 64support Myiobius (formerly Terenotriccus) eryth-rurus as sister group to the other Myiobius spe-cies. This is congruent with Lanyon’s (1988b)merger of erythrurus into Myiobius. Characters40, 48, and, in some most-parsimonious trees,22 support Pyrrhomyias and Hirundinea as sistergroups in agreement with Lanyon (1986,1988b). Neopipo is supported as their sistergroup by characters 11, 43, and 54. This agreeswith one of Mobley and Prum’s (1995) hypoth-eses. However, cranial data for Neopipo are stillneeded to evaluate this hypothesis.

The nasal septa of ’’Myiophobus’’ (lintoni, och-raceiventris, phoenicomitra, and roraimae) havetrabecular plates that are elevated above thebase of the septum (character 4, state 2) whichcaused Lanyon (1986, 1988a) to remove them

from Myiophobus and place them in the Elaeniaassemblage. He refrained from creating newgeneric names until more comparative data areavailable. These results do not give strong sup-port for the removal of lintoni and ochraceiven-tris. Myiophobus, now including only the speciescryptoxanthus, fasciatus, flavicans, inornatus, andpulcher, is supported as the sister group to’’Myiophobus’’ lintoni and ochraceiventris insome most-parsimonious trees by character 49and, additionally in some trees by 17. However,in no most-parsimonious trees are ’’Myioph-obus’’ phoenicomitra and roraimae closely relatedto other Myiophobus. These results support Lan-yon’s (1986, 1988a) conclusion that Myiophobusas traditionally delimited (Traylor 1979, Sibleyand Monroe 1990) is not monophyletic, with atleast phoenicomitra and roraimae belonging else-where. In all most-parsimonious trees, ’’Myio-phobus’’ phoenicomitra and roraimae are the sistergroup of a clade composed of Machetornis andMuscigralla. The relationship is supported bycharacters 8, 9, or both. The interorbital septum(character 9) requires further study in thesetaxa, particularly in Machetornis for which I fol-lowed Warter (1965) and coded as having twofenestrae open (state 0). The one specimen ofMachetornis examined (AMNH 6657), however,had the infraorbital fenestra enlarged and su-praorbital fenestra reduced (state 1), suggest-ing some unknown degree of variation.

Myiarchus assemblage. The monophyly ofLanyon’s (1985) Myiarchus assemblage is sup-ported in some, but not all, most-parsimonioustrees. This clade is supported by the presenceof cavity nesting (character 62). A restricted as-semblage composed only of Rhytipterna, Casior-nis, Sirystes, and Myiarchus is supported in allmost-parsimonious trees by the presence of in-ternal cartilages that are robust ‘‘J’’ or ‘‘L’’shaped (character 32) and are attached to theventral side of the tracheobronchial junction(character 39). In some most-parsimonioustrees it is additionally supported by an ossifiedinterorbital septum (character 9). A Deltarhyn-chus–Ramphotrigon clade is supported in allmost-parsimonious trees by characters 37 and49, and, in some most-parsimonious trees, ad-ditionally by character 11. Lanyon (1985) de-scribed extreme similarity among the craniaand syringes of Myiarchus, Sirystes, and Casior-nis. He was unable to develop a hypothesis oftheir relationships. My results suggest that


Sirystes and Myiarchus are sister groups. Thisrelationship is supported by the presence ofdorsally divergent B1 and B2 syringeal supportelements (character 21), a character that Lanyon(1984) used in the kingbird assemblage but ap-parently did not recognize in the myiarchines.

Elaenia assemblage. These results do notsupport monophyly of the Elaenia assemblage(Lanyon 1988a). Seven extra steps are requiredto force its monophyly, but such trees are notsignificantly different from most-parsimonioustrees (P . 0.19; 14 , n , 25 characters requir-ing differing numbers of steps on the trees be-ing compared). Some most-parsimonious treesinclude a large clade containing most of thegenera of the Elaenia assemblage plus the re-stricted Empidonax assemblage (Fig. 6). In theremainder of most-parsimonious trees thisclade also includes a group of ten taxa (Myio-triccus through Muscigralla in Fig. 5), most ofwhich Lanyon (1988a) placed in his Elaenia as-semblage. In all most-parsimonious trees, thefusion of tracheal A elements into a drum(character 14) supports a ‘‘tyrannulet’’ group(Fig. 6) corresponding roughly to Lanyon’s(1988a) Elaenia group in his Elaenia assemblage.Its monophyly is not supported in the strictconsensus (Fig. 2) because Inezia, Pseudocolop-teryx, and Polystictus are included in the groupin some most-parsimonious trees. That groupis characterized in all most-parsimonious treesby the apparently homologous presence ofwell-developed Mm. obliqui ventrales (character44), a superciliary eye stripe (character 55),perch-gleaning foraging behavior (character61), and unmarked eggs (character 68).

These results support the monophyly of sev-eral of Lanyon’s (1988a) smaller clades withinthe Elaenia assemblage. A clade composed ofElaenia, Myiopagis, and Tyrannulus is supportedby character 14 plus either 46 or 55. Sister-group relationships between Stigmatura andPseudelaenia (character 6), between Pseudotric-cus and Corythopis (characters 5, 18, 33), be-tween Serpophaga and Anairetes (character 14plus either 7 or 43), and between Mionectes andLeptopogon (characters 6, 42, 66, and, in somemost-parsimonious trees, 68) are supported.Tyranniscus is supported as the sister group to’’Mecocerculus’’ calopterus and minor by charac-ters 14 and 35 and, in some most-parsimonioustrees, character 6. This supports Lanyon’s(1988a) grouping of these taxa and his conclu-

sion that the genus Mecocerculus as traditionallydelimited (Traylor 1979, Sibley and Monroe1990) is polyphyletic. In some most-parsimo-nious trees, ’’Mecocerculus’’ (hellmayri, poecilo-cercus, and stictopterus) is the sister group to theclade composed of Tyranniscus and ’’Mecoce-rculus’’ calopterus and minor based on the pres-ence of an additional, posterior trabecular platein the nasal septum (character 6). Lanyon(1988a) left one species (leucophrys) in Mecocer-culus. In no most-parsimonious trees is it close-ly related to the other species traditionallyplaced in Mecocerculus.

Additional tyrannid genera. This analysis re-sulted in hypotheses of relationships for mostof the problematic genera that Lanyon (1986,1988a) maintained incertae sedis. A clade com-posed of Tachuris, Phyllomyias fasciatus and P.griseiceps is supported by the loss of two de-rived characters: the elevated trabecular platein the nasal septum (character 4; cranial datafor P. griseiceps lacking) and a Myiarchus-likeconfiguration of the syringeal B1 and B2 ele-ments (character 22). They also share a derivedsyringeal character with Stigmatura and Pseu-delaenia: a cartilaginous bronchial plate with ro-bust internal cartilages (character 27) whichsupports a clade composed of these five taxa.Lanyon (1988a) thought the syringes ofPhyllomyias fasciatus and P. griseiceps to be verydifferent from each other and speculated thatPhyllomyias (composed of fasciatus, griseiceps,and griseocapilla) is polyphyletic. I examinedLanyon’s cleared-and-stained syringeal speci-mens (three of fasciatus and one of griseiceps)and found them to be quite similar. The syrinxof griseiceps gives the impression of being amore ossified version of the fasciatus syringes.Their cartilaginous bronchial plates are verysimilar in detail and the internal cartilages ofboth species are attached to narrow caudal ex-tensions of the bronchial plates. Besides the de-gree of ossification, the only conspicuous dif-ference between the syringes of the two speciesis the shorter internal cartilages of griseicepswhich lack the large, amorphous ventral andcaudal extension present in fasciatus. Phyllom-yias fasciatus and P. griseiceps are supported assister groups in all most-parsimonious trees bycharacter 61 (fruit/upward hover-glean forag-ing behavior). I suggest that Phyllomyias fascia-tus and P. griseiceps are indeed closely related,possibly sister species (pending examination of


crania of P. griseiceps and P. griseocapilla and sy-ringes of P. griseocapilla).

Machetornis and Muscigralla are supported assister groups by three characters: the loss of aMyiarchus-like configuration of the syringealB1 and B2 elements (character 22), habitation ofopen grassland or marsh (character 60), andground specialist foraging behavior (character61). Colonia and Sublegatus are supported as sis-ter groups by character 55 and, in some most-parsimonious trees, additionally by characters9, 30, and 46.

Homology of nasal septum characters. My re-sults provide a test of the homology of each ofthe states of the nasal septum that Lanyon(1984, 1985, 1986, 1988a, c) used to define hisassemblages. Lanyon (1988c) used the unossi-fied nasal septum (character 1) as one of threecharacters defining his flatbill and tody-tyrantassemblage. When reconstructed on all most-parsimonious trees, the unossified septum ishomologous in Cnipodectes, Onychorhynchus, To-dirostrum, Poecilotriccus, Hemitriccus, Lophotric-cus, and Oncostoma, and, in some most-parsi-monious trees, Rhynchocyclus, Tolmomyias, andPlatyrinchus as well. The unossified septum inTachuris, which Lanyon (1988c) did not includein his flatbill and tody-tyrant assemblage, isconvergent in all most-parsimonious trees. Theunossified septum is symplesiomorphic in allmost-parsimonious trees because, except forthe highly ossified nasal capsules of Dendroco-laptes, furnarioids have unossified septa. Thatcharacter, therefore, does not provide a syna-pomorphy for a flatbill and tody-tyrant assem-blage. However, informative characters are po-tentially existent in the morphology of thecartilaginous parts of the nasal capsules ofthese birds. At least some of the tody-tyrantsprobably have cartilaginous trabecular plateswithin their cartilaginous septa as suggestedby the ossification of those structures in occa-sional specimens (Lanyon 1988c).

Lanyon (1984) defined his kingbird assem-blage partly by the presence of an ossified nasalseptum that lacks both a trabecular plate andan internal support rod. In all most-parsimo-nious trees, a reduced internal support rod(character 3) is homologous in all kingbird as-semblage taxa. In all most-parsimonious trees,character 3 is convergent in Attila, in Colonia,and in Machetornis. The absence of a trabecularplate (character 4) is symplesiomorphic in all

most-parsimonious trees and does not providea synapomorphy of the kingbird assemblage.

Lanyon (1985) defined his Myiarchus assem-blage (without Attila) partly by the presence ofan ossified nasal septum with a conspicuousinternal support rod (character 3). Most tyran-nids possess this character. My family-wideanalysis indicates that it is symplesiomorphicand does not provide a synapomorphy of theMyiarchus assemblage. My results indicate thatthere are no nasal septum characters support-ing this group, with or without Attila. Lanyon(1985) recognized that these genera (except At-tila) have ossified nasal capsules (character 8)which surround the septum, but he did not ex-plicitly include this in his list of characters. Inall most-parsimonious trees in the presentanalysis, character 8 is homologous in Del-tarhynchus, Ramphotrigon, Rhytipterna, Sirystes,Casiornis, and Myiarchus, though in some most-parsimonious trees it is not a synapomorphy. Iexamined the same four crania of Attila spadi-ceus and two of A. cinnamomeus that Lanyon(1985) did and, contrary to his report, foundone (A. cinnamomeus, AMNH 11616) with acompletely ossified nasal capsule. A largersample of Attila crania may show a significanttendency for nasal capsules to ossify and thusmay strengthen the evidence of a relationshipbetween Attila and the Myiarchus assemblage.

Lanyon (1986) defined his Empidonax assem-blage solely by the presence of an ossified nasalseptum with basal trabecular plate and ante-rior notch (character 4) and posterior forking(character 7). In all most-parsimonious trees, abasal trabecular plate and anterior notch is ho-mologous in all genera in the restricted Empi-donax assemblage (Fig. 4). In all most-parsi-monious trees, this character is convergent inMyiobius and in the Pyrrhomyias–Hirundineaclade. This type of nasal septum is also presentin Myiophobus (composed of cryptoxanthus, fas-ciatus, flavicans, inornatus and pulcher) and is, insome most-parsimonious trees, hypothesizedto be homologous with those in Pyrrhomyiasand Hirundinea. Lanyon (1988a) described thesurprising appearance of this type of nasal sep-tum in six species of Elaenia, providing furtherevidence that this seemingly unique characterhas arisen convergently. In these Elaenia spe-cies and in Pyrrhomyias, Hirundinea, and Myio-bius, the trabecular plate is not conspicuouslyforked posteriorly (character 7) as it is in the re-


stricted Empidonax assemblage and in Myiop-hobus. In all most-parsimonious trees, a wideposterior fork is homologous in all genera inthe restricted Empidonax assemblage (exceptlost in Cnemarchus). In all most-parsimonioustrees it is convergent in Myiophobus. Nasal septawith basal trabecular plates and anterior notch-es (character 4) and wide posterior forking(character 7) appear to be synapomorphies of arestricted Empidonax assemblage.

Lanyon (1988a) defined his Elaenia assem-blage solely by the presence of an ossified nasalseptum with a trabecular plate elevated abovethe ventral edge of the septum (character 4). Inall most-parsimonious trees, an elevated tra-becular plate is homologous in all genera of‘‘tyrannulets’’ and allies (Pseudotriccus throughTyranniscus in Fig. 6). It is also homologouswithin a smaller group (Myiotriccus throughMuscigralla in Fig. 5). In all most-parsimonioustrees, the elevated trabecular plate is a syna-pomorphy of a clade containing some or allElaenia assemblage genera and the restrictedEmpidonax assemblage genera. In all most-par-simonious trees, the basal plates of the restrict-ed Empidonax assemblage are hypothesized tohave evolved from elevated plates. In all most-parsimonious trees, the narrow, slender poste-rior forks (character 7) found in several Elaeniaassemblage genera arise two or three indepen-dent times, but are always homologous in Cap-siempis, Phaeomyias, and Nesotriccus, and in Ser-pophaga and Anairetes, and in Pseudocolopteryxand Polystictus. Lanyon (1988a) examined onecranium each of Phyllomyias fasciatus and Tachu-ris and both lacked trabecular plates (the sep-tum of Tachuris was simply unossified). In allmost-parsimonious trees, these are recon-structed as a single derived loss of ossified, el-evated trabecular plates.

Levels of homoplasy in character suites. Cra-nial characters (mean RC 5 0.39), syringealcharacters (mean RC 5 0.47), plumage charac-ters (mean RC 5 0.33), and behavioral and eco-logical characters (mean RC 5 0.52) did not dif-fer significantly in levels of homoplasy (P 50.57). The behavioral and ecological charactersin this study contained a level of phylogeneticinformation similar to those that Hughes (1996)used for the Cuculidae. These results are in ac-cord with a growing number of studies thatfind significant phylogenetic structure in be-havioral and ecological characters (e.g. Prum

1990a, Winkler and Sheldon 1993, Paterson etal. 1995, Hughes 1996, McLennan and Mattern2001). Among the suites of morphological char-acters, plumage characters exhibited the mosthomoplasy, though not significantly so. Onlyone plumage character—long rictal bristles(character 57)—exhibited no homoplasy on anymost-parsimonious tree. A pointed crest (char-acter 58) exhibited no homoplasy on some, butnot all most-parsimonious trees. The ten plum-age characters that describe color patterns tendto be more homoplasious (mean RC 5 0.26)than those that describe feather form, but thisis still not significantly different from othercharacter suites (P 5 0.36). Omland and Lan-yon (2000) found similarly high levels of ho-moplasy in plumage characters in orioles andcautioned against heavy use of such charactersin phylogenetic analysis. It appears likely thatconvergence, parallelism, and reversal havebeen common in the evolution of avian plum-age color patterns, perhaps due to evolutionaryconservatism of the underlying developmentalpathways coupled with lability of the on–offswitches of those pathways (Price and Pavelka1996).

Summary. I performed a cladistic analysisof all currently available morphological and be-havioral data for the Tyrannidae. Most charac-ters were drawn from Lanyon (1984, 1985,1986, 1988a, b, c). My analysis differs from Lan-yon’s in five important ways: first, I added sev-eral characters. Second, I examined furnarioidoutgroups in addition to the tyrannoid out-groups Lanyon considered. Third, I assessedcharacter states across all tyrannid genera andin outgroups, allowing simultaneous analysisof all taxa and tests of monophyly of the Tyran-nidae and of each of Lanyon’s proposed assem-blages. Fourth, I gave characters equal weight,allowing tests of the homology of cranial andsyringeal characters that Lanyon used to definegroups. Fifth, I worked within a cladisticframework that explicitly roots trees to an out-group. This ensures plesiomorphies are notused as character support for clades.

My results provide equivocal support formonophyly of the Tyrannidae. An equally par-simonious hypothesis suggests that the flatbilland tody-tyrant genera are more distantly re-lated to other tyrannids than are cotingids andpiprids. Results support monophyly of thekingbird assemblage, restricted Myiarchus and


Empidonax assemblages, and several lower-lev-el groupings. Monophyly of neither the Elaenianor the flatbill and tody-tyrant assemblage issupported. Nasal septum characters that pastworkers have considered conservative and phy-logenetically informative are supported assynapomorphies of the kingbird assemblageand of a restricted Empidonax assemblage. Anasal capsule character provides equivocalsupport for a Myiarchus assemblage withoutAttila. Many nodes have minimal support.Much more data are needed to resolve relation-ships in the Tyrannidae, especially at higherlevels. In accord with previous studies, plum-age coloration patterns exhibited relativelyhigh levels of homoplasy.

ACKNOWLEDGMENTS

This paper is based on a dissertation submitted inpartial fulfillment of the requirements for a Ph.D. inthe Department of Zoology, Michigan State Univer-sity. I thank D. Straney, R. Prum, W. Lanyon, D.Brooks, R. Leschen, M. Donoghue, J. White, and T.Getty for helpful discussion and encouragement.This paper benefited from comments by B. Livezey,R. Zink, and an anonymous reviewer. I thank G. Bar-rowclough for access to the collections at the Amer-ican Museum of Natural History. This work was sup-ported by the W. K. Kellogg Biological StationGraduate Research Training Group (RTG) funded byNSF DIR-09113598, by a Michigan State UniversityCollege of Natural Science Doctoral Recruiting Fel-lowship, by the Frank M. Chapman Fund of theAmerican Museum of Natural History, and by aGrant-in-Aid-of-Research from Sigma Xi. This iscontribution no. 868 of the W. K. Kellogg BiologicalStation.

LITERATURE CITED

ADAMS, E. N. 1972. Consensus techniques and thecomparison of taxonomic trees. Systematic Zo-ology 21:390–397.

AMERICAN ORNITHOLOGISTS’ UNION. 1998. Check-list of North American Birds, 7th ed. AmericanOrnithologists’ Union, Washington, D.C.

AMES, P. L. 1971. The morphology of the syrinx inpasserine birds. Bulletin of the Peabody Muse-um of Natural History, no. 37.

BAUMEL, J. J. 1979. Osteologia. Pages 53–121 in No-mina Anatomica Avium (J. J. Baumel, A. S. King,A. M. Lucas, J. E. Breazile, and H. E. Evans,Eds.). Academic Press, London.

BOSTWICK, K. S. 2000. Display behaviors, mechanicalsounds, and evolutionary relationships of the

Club-winged Manakin (Machaeropterus delicio-sus). Auk 117:465–478.

BREMER, K. 1988. The limits of amino acid sequencedata in angiosperm phylogenetic reconstruction.Evolution 42:795–803.

DEQUEIROZ, A., AND P. H. WIMBERGER. 1993. Theusefulness of behavior for phylogeny estima-tion—levels of homoplasy in behavioral andmorphological characters. Evolution 47:46–60.

FELSENSTEIN, J. 1982. Numerical methods for infer-ring evolutionary trees. Quarterly Review of Bi-ology 57:379–404.

FITZPATRICK, J. W. 1980. Foraging behavior of Neo-tropical tyrant flycatchers. Condor 82:43–57.

GRIFFITHS, C. S. 1994. Monophyly of the Falconifor-mes based on syringeal morphology. Auk 111:787–805.

HILTY, S. L., AND W. L. BROWN. 1986. A Guide to theBirds of Colombia. Princeton University Press,Princeton, New Jersey.

HUGHES, J. M. 1996. Phylogenetic analysis of the Cu-culidae (Aves: Cuculiformes) using behavioraland ecological characters. Auk 113:10–22.

KALLERSJO, M., J. S. FARRIS, A. G. KLUGE, AND C.BULT. 1992. Skewness and permutation. Cladis-tics 8:275–288.

KING, A. S. 1979. Systema respiratorium. Pages 227–265 in Nomina Anatomica Avium (J. J. Baumel,A. S. King, A. M. Lucas, J. E. Breazile, and H. E.Evans, Eds.). Academic Press, London.

LANYON, S. M. 1985. Molecular perspective on high-er-level relationships in the Tyrannoidea (Aves).Systematic Zoology 34:404–418.

LANYON, W. E. 1984. A phylogeny of the kingbirdsand their allies. American Museum Novitates,no. 2797.

LANYON, W. E. 1985. A phylogeny of the myiarchineflycatchers. Ornithological Monographs 36:361–380.

LANYON, W. E. 1986. A phylogeny of the thirty-threegenera in the Empidonax assemblage of tyrantflycatchers. American Museum Novitates, no.2846.

LANYON, W. E. 1988a. A phylogeny of the thirty-twogenera in the Elaenia assemblage of tyrant fly-catchers. American Museum Novitates, no. 2914.

LANYON, W. E. 1988b. The phylogenetic affinities ofthe flycatcher genera Myiobius Darwin and Te-renotriccus Ridgeway. American Museum Novi-tates, no. 2915.

LANYON, W. E. 1988c. A phylogeny of the flatbill andtody-tyrant assemblage of tyrant flycatchers.American Museum Novitates, no. 2923.

LEE, P. L. M., D. H. CLAYTON, R. GRIFFITHS, AND R.D. M. PAGE. 1996. Does behavior reflect phylog-eny in swiftlets (Aves: Apodidae)? A test usingcytochrome b mitochondrial DNA sequences.Proceedings of the National Academy of Scienc-es USA 93:7091–7096.


MADDISON, W. P., AND D. R. MADDISON. 1992.MacClade: Analysis of Phylogeny and CharacterEvolution, version 3.04. Sinauer Associates, Sun-derland, Massachusetts.

MARINI, M. A. 1992. Foraging behavior and diet ofthe Helmeted Manakin. Condor 94:151–158.

MCKITRICK, M. C. 1985. Monophyly of the Tyranni-dae (Aves): Comparison of morphology andDNA. Systematic Zoology 34:35–45.

MCLENNAN, D. A., AND M. Y. MATTERN. 2001. Thephylogeny of Gasterosteidae: combining behav-ioral and morphological data sets. Cladistics 17:11–27.

MOBLEY, J. A., AND R. O. PRUM. 1995. Phylogeneticrelationships of the Cinnamon Tyrant, Neopipocinnamomea, to the tyrant flycatchers (Tyranni-dae). Condor 97:650–662.

OMLAND, K. E., AND S. M. LANYON. 2000. Recon-structing plumage evolution in orioles (Icterus):Repeated convergence and reversal in patterns.Evolution 54:2119–2133.

PATERSON, A. M., G. P. WALLIS, AND R. D. GRAY.1995. Penguins, petrels, and parsimony: Doescladistic analysis of behavior reflect seabirdphylogeny? Evolution 49:974–989.

PRICE, T., AND M. PAVELKA. 1996. Evolution of a col-our pattern: History, development, and selec-tion. Journal of Evolutionary Biology 9:451–470.

PRUM, R. O. 1990a. Phylogenetic analysis of the evo-lution of display behavior in the Neotropicalmanakins (Aves: Pipridae). Ethology 84:202–231.

PRUM, R. O. 1990b. A test of the monophyly of themanakins (Pipridae) and of the cotingas (Cotin-gidae) based on morphology. Occasional Papersof the Museum of Zoology, University of Mich-igan, no. 723.

PRUM, R. O. 1992. Syringeal morphology, phylogeny,and evolution of the Neotropical manakins(Aves: Pipridae). American Museum Novitates,no. 3043.

PRUM, R. O. 1994. Phylogenetic analysis of the evo-lution of alternative social behavior in the man-akins (Aves: Pipridae). Evolution 48:1657–1675.

PRUM, R. O. 1997. Phylogenetic tests of alternative in-tersexual selection mechanisms: Trait macro-evolution in a polygynous clade (Aves: Pipri-dae). American Naturalist 149:668–692.

PRUM, R. O., AND W. E. LANYON. 1989. Monophylyand phylogeny of the Schiffornis group (Tyran-noidea). Condor 91:444–461.

RIDGELY, R. S., AND G. TUDOR. 1994. The Birds ofSouth America, vol. 2. The Suboscine Passerines.University of Texas Press, Austin.

SIBLEY, C. G., AND J. E. AHLQUIST. 1985. Phylogenyand classification of New World suboscine pas-serine birds (Passeriformes: Oligomyodi: Tyran-nides). Ornithological Monographs 36:396–428.

SIBLEY, C. G., AND B. L. MONROE, JR. 1990. Distri-bution and Taxonomy of Birds of the World. YaleUniversity Press, New Haven, Connecticut.

SKUTCH, A. F. 1960. Life histories of Central Ameri-can birds II. Cooper Ornithological Society,Berkeley, California.

SNOW, D. W. 1982. The Cotingas. Cornell UniversityPress, Ithaca, New York.

SORENSON, M. D. 1996. TREEROT. University ofMichigan, Ann Arbor.

STILES, F. G., AND A. F. SKUTCH. 1989. A Guide to theBirds of Costa Rica. Cornell University Press,Ithaca, New York.

SWOFFORD, D. L. 1991. When are phylogeny esti-mates from molecular and morphological dataincongruent? Pages 295–333 in PhylogeneticAnalysis of DNA Sequences (M.M. Miyamotoand J. Cracraft, Eds.). Oxford University Press,New York.

SWOFFORD, D. L. 2000. PAUP*. Phylogenetic AnalysisUsing Parsimony (*and Other Methods), version4. Sinauer Associates, Sunderland, Massachusetts.

TEMPLETON, A. R. 1983. Phylogenetic inference fromrestriction endonuclease cleavage site maps withparticular reference to the evolution of humansand the apes. Evolution 37:221–244.

TRAYLOR, M. A., JR. 1977. A classification of the ty-rant flycatchers (Tyrannidae). Bulletin of theMuseum of Comparative Zoology 148:129–184.

TRAYLOR, M. A., JR. 1979. Family Tyrannidae. Pages1–245 in Check-list of Birds of the World, vol.VIII (M. A. Traylor, Jr., Ed.). Museum of Com-parative Zoology, Cambridge, Massachusetts.

TRAYLOR, M. A., JR., AND J. W. FITZPATRICK. 1982. Asurvey of the tyrant flycatchers. Living Bird 19:7–50.

WARTER, S. L. 1965. The cranial osteology of the NewWorld Tyrannoidea and its taxonomic implica-tions. Ph.D. dissertation, Louisiana State Uni-versity, Baton Rouge.

WINKLER, D. W., AND F. H. SHELDON. 1993. Evolutionof nest construction in swallows (Hirundinidae):A molecular phylogenetic perspective. Proceed-ings of the National Academy of Sciences USA90:5705–5707.

Associate Editor: R. Zink

APPENDIX 1. Descriptions of characters used in anal-ysis. Characters 6, 11, and 13 are ordered. Character11 is ordered by a character state tree described inFigure 1. The remaining characters are unordered.

Cranial

Nomenclature follows Baumel (1979) when possi-ble. Otherwise, it follows Lanyon (1984, 1985, 1986,1988a, b, c).


1. Septum nasale.—(0) Ossified to varying degrees;(1) unossified in the flatbill and tody-tyrant assem-blage (Lanyon 1988c), in Tachuris (Lanyon 1988a),and in most furnarioids (pers. obs.). In these taxathere is usually just a shallow dorsal remnant of aseptum in cleaned museum specimens.

2. Anterior segment of septum nasale osseum.—(0)Well ossified anterior to the internal support rod; (1)poorly ossified anterior to support rod.

3. Internal support rod.—(0) Conspicuous internal sup-port rod within the septum nasale osseum; (1) no con-spicuous internal support rod. In the kingbird assem-blage (Lanyon 1984) and in the genera Attila, Colonia,and Machetornis (J. S. Birdsley pers. obs.), there is onlya pattern or grain running anterodorsally to postero-ventrally on the surface of the septum in the positionwhere the rod is located in the septa of othertyrannoids.

4. Transverse trabecular plate within septum nasal os-seum.—(0) No transverse trabecular plate; (1) a trans-verse trabecular plate is at the base (ventral edge) ofthe septum and there is a distinctive notch in the sep-tum anterior to the trabecular plate; (2) a transversetrabecular plate is elevated above the ventral edge ofthe septum, so that there appears to be a mid-sagittalridge on the ventral side of the trabecular plate.There is no anterior notch associated with this typeof trabecular plate; (3) a transverse trabecular plateis barely elevated above the ventral edge of the sep-tum and occupies only a very short segment of theanterior part of the septum. It is very broad, trun-cated posteriorly, and has no anterior notch. Presentin Colonia and hypothesized by Lanyon (1986) to beunique among tyrannids.

5. Trabecular plate reduced, anterior.—(0) The elevat-ed type trabecular plate (if present) is relativelybroad and occupies the length of the septum anteriorto the internal support rod; (1) the elevated trabec-ular plate is narrow and occupies only a small partof the most anterior section of the septum.

6. Trabecular plate divided.—(0) The trabecular plate(if present) exists only anterior to the internal sup-port rod; (1) the elevated type trabecular plate is di-vided into a small section posterior to the internalsupport rod and larger section anterior to the rod.Both sections are relatively slim; (2) the elevated tra-becular plate is divided into a bulbous posterior sec-tion and a thin anterior section; (3) the elevated tra-becular plate is divided into bulbous posterior andbulbous anterior sections.

7. Trabecular plate forked posteriorly.—(0) Trabecularplate (if present) is truncate or tapered posteriorly;(1) trabecular plate is widely forked posteriorly (Lan-yon 1986, 1988b). (2) trabecular plate is narrowlyforked posteriorly, appearing as a slender tuningfork (Lanyon 1988a).

8. Nasal capsule.—(0) Unossified; (1) fully ossifiedincluding alinasal walls and concha nasalis [turbi-nals in Lanyon (1985)].

9. Septum interorbitale.—(0) Two open fenestrae; (1)infraorbital fenestra enlarged and supraorbital fe-nestra reduced (Lanyon 1988a, c); (2) both fenestraereduced or absent; septum ossified.

10. Medial ridge in frontal region of cranium.—(0) Ab-sent; (1) present.

Syringeal

Nomenclature of syringeal support elements andcartilages follows Ames (1971) and Lanyon (1984,1985, 1986, 1988a, b, c). Nomenclature of syringealmusculature follows King (1979) which is also con-sistent with the above references.

11. Double, complete A elements.—(0) None of the(primarily tracheal) ‘‘A’’ series of syringeal supportrings completely encircle the bronchi; (1) one A ele-ment is double (associated with the bronchi ratherthan the trachea) and complete (completely encir-cling each bronchus). They are medially cartilagi-nous; (2) two A elements are double, complete, andmedially cartilaginous; (3) two or more A elementsare double, complete, and medially ossified; (4) oneA element is double, complete, and medially ossified;(5) incomplete A3 elements contribute to support ofeach bronchus through fusion to the complete A2 el-ements; (6) incomplete A2 elements contribute tosupport of each bronchus through fusion to the com-plete A3 elements.

12. Cartilaginous dorsal segments of A1 elements.—(0)A1 elements are nearly fully ossified, sometimeswith small, cartilaginous dorsal tips; (1) A1 elementsterminate dorsally in large cartilaginous segments.

13. Medial segments of A2 elements modified.—(0) Me-dial segments of A2 elements (when double andcomplete) are smooth; (1) cartilaginous medial seg-ments of A2 elements are enlarged and bulbous cau-dally, presumably for the attachment of the internalcartilages (Lanyon 1986); (2) cartilaginous medialsegments of A2 elements are enlarged and bulbousas in state (1) but are disconnected from the remain-ing (ossified) portion of the A2 element.

14. Tracheal ‘‘drum’’.—(0) Tracheal A elements arelargely independent of one another; (1) two or moretracheal A elements are fused to form a trachealdrum. The drum is poorly fused and includes A2 andhigher elements; (2) the drum is robust and includesA2 and higher elements; (3) the drum is robust andincludes A3 and higher elements, but not A2.

15. Nodule on lateral surface of A1 and A2.—(0) Lat-eral surfaces of A1 and A2 are smooth; (1) ossifiednodule on the lateral surface of each A1 and A2 el-ement, with cartilaginous connections betweenthem.

16. Pessulus divides tracheal A elements dorsally.—(0)Tracheal A elements are complete dorsally; (1) ossi-fied pessulus extends anteriorly on the dorsal side ofthe trachea to divide four or more A elements.


17. Pessulus extends from the first single, complete Aelement.—(0) Pessulus (when present) is not attachedto the first single, dorsally complete A element; (1)pessulus is an ossified caudal extension of the dorsalside of the first single, dorsally complete A element.

18. Pessulus large, concave.—(0) Pessulus (when pre-sent) is convex dorsally; (1) the ossified pessulus isrelatively large and concave dorsally. Its greatestbreadth is at the point where the internal cartilagesattach.

19. B1-2 connected ventrally.—(0) The bronchial,cartilaginous B1 and B2 elements are independent attheir ventral tips; (1) B1 and B2 are connected at theirventral tips. The connection is flattened or acute; (2)B1 and B2 are connected at their ventral tips. Theconnection is symmetrically rounded.

20. B1-2 connections are close medially.—(0) The leftand right ventral ends of B1 and B2 are relatively farapart; (1) the bronchi are close together in the regionof B1 and B2, so that the left and right ventral con-nections of those elements nearly touch one anothermedially.

21. B1-2 diverge dorsally.—(0) B1 and B2 are nearlyparallel dorsally; (1) B1 and B2 diverge dorsally sothat their dorsal tips lie relatively far apart.

22. B1-2 shaped as in Myiarchus.—(0) B1 is narrowdorsally or B2 is narrow ventrally; (1) B1 and B2 havea distinct configuration as typified by Myiarchus(Lanyon 1985): B1 is narrow except for a very broaddorsal end. B2 is narrow except for a broad, trian-gular ventral end which is barely attached to the ven-tral end of B1.

23. B1 breadth.—(0) Breadth of the lateral region ofB1 is less than or equal to the breadth at the ventraland dorsal tips; (1) B1 is broader laterally than at thetips.

24. B1 bulbous dorsally.—(0) B1 flat dorsally; (1) B1bulbous or swollen at dorsal tips.

25. B2 Y-shaped ventrally.—(0) Ventral end of B2 isrounded or truncate; (1) ventral end of B2 is forkedor shaped like the letter ‘‘Y’’.

26. Syrinx and trachea laterally compressed.—(0) Tra-chea and tracheobronchial junction are not both lat-erally compressed; (1) in Lophotriccus and Oncostoma(Lanyon 1988c), the entire trachea and tracheobron-chial junction are laterally compressed so that theyare unusually narrow when viewed dorsally or ven-trally, and broad when viewed laterally. Similar mor-phology occurs in Machetornis where the tracheo-bronchial junction, but not the trachea, is laterallycompressed (J. S. Birdsley pers. obs.).

27. Cartilaginous bronchial plate.—(0) Dorsal tips ofthe incomplete bronchial A elements are not con-nected with one another or with the pessulus; (1) ahorseshoe-shaped cartilaginous plate connects thedorsal tips of the incomplete bronchial A elementsand the pessulus. It forms the dorsal or craniodorsalmargins of the medial tympaniform membranes. Theinternal cartilages are very small, delicate rods lo-

cated near the caudal ends of the horseshoe, some-times attached to it; (2) a cartilaginous plate as instate (1), but the internal cartilages are robust.

28. Cartilaginous tracheal plate.—(0) Lower trachealA elements are ossified dorsally; (1) cartilaginouspessulus extends cranially so that the lower trachealA elements up to A6 or A8 are cartilaginous dorsally.

29–39. Internal cartilages.—The internal tympani-form membranes in the syringes of all tyrannids andsome cotingids contain cartilaginous structureswhich are often attached to A elements. Becausethere is extreme variation in the shape, position, andattachment of internal cartilages, Prum and Lanyon(1989) hypothesized them to be derived in cotingidsindependently of those in tyrannids. Prum (1990b)questions their homology even within tyrannids, andrecommends using unique, detailed morphologies ofinternal cartilages as characters, while refrainingfrom the hypothesis of homology of internal carti-lages as broadly defined. Here I follow that recom-mendation and the precedent of Lanyon (1984, 1985,1986, 1988a, b, c) in coding shared, detailed mor-phologies of tyrannid internal cartilages as indepen-dent, derived characters.

29. Narrow, linear.—(0) Absent; (1) cartilages arenarrow and basically linear.

30. Triangular.—(0) The base (the cranial end) isnarrower than or scarcely greater than the width ofthe rest of the cartilage; (1) the cartilage is triangular,with the base much broader than the rest of thecartilage.

31. Spatulate.—(0) Absent; (1) cartilages are broadand shaped like a spatula or child’s mitten with athickened dorsal edge.

32. ‘‘J’’ or ‘‘L’’ shaped.—(0) Absent; (1) cartilages arethin and bent in a ‘‘J’’ or ‘‘L’’ shape; (2) cartilages arerobust and bent in a ‘‘J’’ or ‘‘L’’ shape.

33. Forked.—(0) Absent; (1) cartilages are forkedcaudally.

34. Ventral extension.—(0) Absent; (1) cartilageshave poorly staining, amorphous ventral extensionsfrom their caudal halves; (2) cartilages have well-formed, squarish ventral extensions from their cau-dal halves.

35. Broad, amorphous caudal segments.—(0) Absent;(1) cartilages are narrow cranially but broaden andbecome amorphous caudally.

36. Narrow, curved.—(0) Absent; (1) cartilages arerelatively narrow and curved (medially concave).

37. Straight, expanded caudally.—(0) Absent; (1) car-tilages are relatively long, straight, and expandedcaudally. They are somewhat twisted, with the cra-nial half flattened at right angles to the internaltympaniform membrane and the caudal half flat-tened within the plane of the membrane.

38. Attached to tracheal drum and incomplete A ele-ments.—(0) Cartilages are attached either to an in-complete A element or to a tracheal drum, but notboth; (1) cartilages are attached to the dorsal end of


an incomplete A element (either A1 or A2) as well asto a tracheal drum.

39. Attached to ventral side of tracheobronchial junc-tion.—(0) Cartilages are attached to the dorsal sideof various structures in the tracheobronchial junc-tion; (1) cartilages are attached to ventral or medio-ventral segments of A2 or A3.

40. Narrow strand of cartilage between B2 and B3.—(0) Absent; (1) in addition to the large internal car-tilages, there is a narrow strand of cartilage in theinternal tympaniform membrane uniquely placedbetween the ventral ends of B2 and B3.

41. Cartilaginous plug of tissue between A2 ele-ments.—(0) The space just caudal to the tracheobron-chial junction is vacant; (1) there is a plug of tissuejust caudal to the tracheobronchial junction and be-tween the cartilaginous medial segments of the A2elements. This plug stains weakly for cartilage.

42. Mm. tracheolaterales.—(0) Cover only the lat-eral portions of the trachea; (1) right and left Mm.tracheolaterales widen ventrally and meet aboveA20, sometimes lower, and completely cover the ven-tral and ventrolateral surface of the trachea fromthere posteriad, diverging just before their insertionin the region of the syrinx; (2) Mm. tracheolateralescover the ventral and ventrolateral surface of the tra-chea as in state (1) but do not diverge at their inser-tion. Instead, they converge to a point and insertmidventrally on the lower A elements.

43. Mm. obliqui ventrales.—(0) Intrinsic syringealmuscles (if present) are different from Mm. obliquiventrales in detail of fiber direction and form of in-sertion (Prum and Lanyon 1989; Prum 1990b, 1992);(1) Mm. obliqui ventrales insert on the ventral or lat-eral side of the syrinx (Ames 1971, McKitrick 1985,Mobley and Prum 1995). (2) Mm. obliqui ventrales in-sert on the dorsal side of the syrinx (Ames 1971).

44. Mm. obliqui ventrales well developed.—(0) Mm.obliqui ventrales (if present) are relatively undevel-oped, appearing flat; (1) Mm. obliqui ventrales are welldeveloped, appearing as round, bulging muscle mas-ses. Present in at least some members of the kingbirdand Elaenia assemblages (Lanyon 1984, 1988a) and inHymenops, Polioxolmis, and Cnemarchus (pers. obs.).

45. Mm. obliqui laterales.—(0) Absent; (1) syrinxhas a pair of laterally-originating and laterally-in-serting intrinsic syringeal muscles. In most taxa pos-sessing them, these muscles exist in addition to Mm.obliqui ventrales, but apparently in some tody-tyrants,Mm. obliqui laterales are the only intrinsic syringealmuscles (Ames 1971).

Plumage

46. Crown patch.—(0) Crown plumage is mono-chromatic; (1) crown plumage contains a patch ofwhite, red, orange, or yellow which is usually con-cealed or semiconcealed.

47. Sexually dichromatic plumage.—(0) Sexes havesimilar or identical plumage color and patterns; (1)sexes have markedly different plumage color andpatterns.

48. Wing patterned.—(0) Remiges are similarly col-ored on their proximal and distal halves; (1) remigesare black or fuscous distally, and buff or cinnamon-rufous proximally; (2) remiges as in state (1) exceptthey are white proximally.

49. Wing bars.—(0) Absent; (1) rufous-buff; (2) yel-low; (3) white.

50. Throat plumage.—(0) Unstreaked; (1) darkstreaks against a white background.

51. Outer rectrix pale.—(0) Outer rectrices are sim-ilarly colored on their inner and outer webs; (1) outerwebs of the outer rectrices are pale (white to cinna-mon–rufous), contrasting with darker inner webs.

52. Tail length.—(0) Short to moderate; (1) elongat-ed, forked.

53. Tail rufous.—(0) Tail is various colors but notentirely rufous; (1) entire tail is rufous.

54. Underparts rufous.—(0) Underparts various col-ors but not entirely rufous; (1) underparts entirelyrufous.

55. Superciliary.—(0) No contrasting stripe over orthrough the eye; (1) white, yellow, or cinnamon su-perciliary stripe over or through the eye.

56. Plumage black and white.—(0) Plumage has atleast some color or is solid black, brown, or white;(1) plumage is black and white or brown and white.

57. Rictal bristles.—(0) Short or absent; (1) as longor nearly as long as the bill.

58. Pointed crest.—(0) Feathers of the crown form arounded crest; (1) feathers of the crown form a point-ed, prominent crest.

59. Primaries notched.—(0) Outer primaries notnotched at their tips; (1) outer one, two, or three pri-maries are notched or attenuated in at least the maleof the species.

Behavior and Ecology

60. Habitat.—(0) Inhabit forests and woodlands; (1)inhabit open grassland or marsh; (2) inhabit opencliffs, canyons, and roadcuts.

61. Foraging behavior.—Fitzpatrick (1980) charac-terized the foraging behavior of nearly every tyran-nid genus and recognized 10 ‘‘foraging mode’’ cat-egories in the family. Here I treat each of theseforaging mode categories as alternative unorderedderived states of a single character. I consulted Snow(1982), Hilty and Brown (1986), and Marini (1992) forbehavior of outgroups.

(0) Fruit/upward hover-glean generalists feedheavily on fruits during most or all of the yearbut also upward hover-glean and perch-gleaninsects.

(1) Fruit/hawk generalists combine frugivorywith aerial-hawking of flying insects. They


use these capture techniques in roughly equalproportions.

(2) Outward hover-glean generalists often gleaninsects from foliage surfaces via an outward ordownward ‘‘sally’’ or capture flight. The birdmay slow down and hover while capturing theinsect or snap it up while in direct flight, oftenstriking the foliage and continuing to a newperch. Upward hover-gleaning and aerial-hawking are additional significant elements oftheir foraging repertoires.

(3) Enclosed perch hawk generalists have foragingrepertoires composed of roughly equal propor-tions of aerial-hawking, upward hover-glean-ing, and upward-striking.

(4) Near-ground generalists usually hunt fromperches near the ground and use a wide vari-ety of capture techniques including perch-to-ground and perch-to-water sallies, upwardand outward hover-gleaning and striking, andaerial-hawking.

(5) Perch-glean specialists capture most of theirprey while perched, without sallying. Hover-gleaning and frugivory are minor componentsof their repertoire.

(6) Upward-strike specialists capture most of theirprey in an explosively rapid upward sally tothe underside of a leaf, snatching or scoopingan insect off the under-surface, usually withouthovering.

(7) Aerial-hawk specialists capture most of theirprey out of the air.

(8) Perch-to-ground specialists hunt from lowperches, usually sallying to the ground tocatch prey. Aerial-hawking may be a minorcomponent of their repertoire.

(9) Ground specialists stand, walk, or run on theground and search for their prey which theypick off the ground or low vegetation. Aerial-hawking from the ground may be a minor com-ponent of their repertoire.

62. Cavity nesting.—(0) Nests are in open situa-tions; (1) nests are located in tree cavities or holes inearthen banks.

63. Enclosed nest within hanging vegetation.—(0)Nest, if hanging, is built around a branch and sus-pended from it; (1) enclosed, globular nest with aside entrance is built within a hanging mass ofvegetation.

64. Enclosed nest pendant.—(0) Nest is not suspend-ed from a branch; (1) enclosed, globular nest with aside or bottom entrance is suspended from a branch;(2) enclosed, elongate (1 m or more) nest with a sideentrance is suspended from a branch.

65. Enclosed, pendant nest retort-shaped.—(0) Nest, ifhanging, has an entrance hole at or above the bottomof the nest chamber; (1) enclosed, pendant, globularnest shaped like a chemists’ retort with a downward-pointing entrance spout.

66. Enclosed nest with visor over entrance.—(0) Nest,if enclosed, has an unsheltered entrance hole; (1) en-closed nest has a short roof or visor over the sideentrance.

67. Nest used as dormitory.—(0) Nests are used onlyduring breeding; (1) adults sleep in their nests at allseasons. Presumed present only in Rhynchocyclus andTolmomyias (Skutch 1960).

Eggs

68. Eggs.—(0) Marked with dark blotches andstreaks; (1) plainly colored, usually some shade ofwhite.


APPENDIX 2. Character-state matrix of 68 characters for 103 tyrannid taxa plus 3 outgroup taxa used inphylogenetic analysis. Asterisks (*) denote character stage polymorphisms of the type ‘‘0/1’’. Other poly-morphic combinations are coded with a slash (e.g. ‘‘0/2’’). Unknown character states are coded ‘‘?’’. SeeAppendix 1 for descriptions of characters.


APPENDIX 2. Continued.

Myiophobus, l,o 5 species lintoni and ochraceiventris. Myiophobus p,r 5 species phoenicomitra and roraimae. Myiophobus 5 species cryptoxanthus,fasciatus, flavicans, inornatus, and pulcher. Mecocerculus c,m 5 species calopterus and minor. Mecocerculus h,p,s 5 species hellmayri, poecilocercus, andstictopterus. Mecocerculus 5 species leucophrys. Myiobius a,b,v 5 species atricaudus, barbatus, and villosus.

PHYLOGENY OF THE TYRANT FLYCATCHERS (TYRANNIDAE) BASED ON

Documents

Transcript of PHYLOGENY OF THE TYRANT FLYCATCHERS (TYRANNIDAE) BASED ON