The Condor 9 I x444-46 I 0 The Cooper Ornithological Society 1989 MONOPHYLY AND PHYLOGENY OF THE SCHIFFORNIS GROUP (TYRANNOIDEA)’ RICHARD 0. PRUM Museum of Zoology and Department of Biology, The Universityof Michigan, Ann Arbor, MI 48109 WESLEY E. LANYON~ Department of Ornithology,The American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024 Abstract. A cladistic analysisof 19 characterswas used to investigate the phylogeny of the Schzjornisgroup, a monophyletic assemblageof six generaof suboscinepasserines which are currently placed in three different tyrannoid families: Schzfirnis (Pipridae, manakins), Laniisoma, Zodopleura(Cotingidae, cotingas), Laniocera, Xenopsaris,and Pachyramphus (Tyrannidae, tyrant flycatchers).The charactersystems analyzed include syringealand cranial morphology, plumage, and nest architecture.Monophyly of the Schijornis groupwas sup- ported by two syringeal synapomorphies.Within the group, the phylogeneticrelationships of five of the six genera were resolved; Schzfirnis is the sister group to Laniisoma and Laniocera, and Xenopsarisis the sister group of Pachyramphus.The relationship of Zodo- pleura to these two corroboratedclades was not resolved. The higher-level relationshipsof the Schzjornis group to other tyrannoids was not resolved. Results of two previous bio- chemical phylogeniesof tyrannoids both corroborate and conflict with this morphological hypothesis.Comparative analysis of the evolution of breeding systemsin tyrannoids indi- cates that polygyny may have developed independently within the group, resulting in the evolution of a novel form a male advertisement behavior without conspicuoussexual di- morphism and a distinct type of spatial dispersion of male territories. Key words: Phylogeneticsystematics; morphology; syrinx; cotingas;manakins; tyrant flycatchers;breeding system evolution; Schiffomis; Pachyramphus. Resumen. Un analisis cladistico de 19 caracteristicasse emplea para investigar las re- lacidnes filogeneticasde1 grupo Schzfirnis, un conjunto monofilttico de seis generos, 10s cualesse ubican ahora en tres familias diferentesen la superfamilia Tyrannoidea: Schzfirnis (Pipridae), Laniisoma, Zodopleura(Cotingidae), Laniocera, Xenopsaris,y Pachyramphus (Tyrannidae). Las caracteristicasempleadas incluyen la morfologia de la siringey de1craneo, el plumaje, y la estructurade1 nido. La posici6n de1grupo Schzjornis en Tyrannoidea no se ha determinado. Dentro de1 grupo, las relaciones de cinco de 10s seis gtneros fueron resueltaspor la hipBtesisfilogenetica mas parsimoniosa, es decir, aquella que requiere el menor numero de caracteristicasderivados. Schzfirnis es m&s cercano a un linaje que consiste de Laniisoma y Laniocera; y Xenopsarises m8s cercano a Pachyramphus.Las relaciones de Zodopleurano eran resolvado. La comparaci6n de estos resultadoscon dos hip6tesesfilogeneticas bioquimicas resultasimilitudes y conflictos.Un analisiscomparative de la evoluci6n de 10scomportamientos reproductivosen Tyrannoidea indica que polyginia se ha desarolladoindependientemente en el grupo Schzfirnis. TambiCn en el group0 Schif- fornis, se han desarollado nuevos tipos de dispersion territorial y anuncio nuptial de 10s machos sin dimorfismo sexual. INTRODUCTION posed of over 500 specieswhich are placed in The New World suboscine superfamily Tyran- the tyrant flycatchers (Tyrannidae), manakins noidea (Passeriformes)is a diverse group com- (Pipridae), cotingas (Cotingidae), plantcutters (Phytotomidae), and sharpbills (Oxyruncidae) I Received 6 October 1988. Final acceptance6 Feb- (Traylor 1979). The phylogenetic interrelation- ruary 1989. ships of the tyrannoids have recently received a * Presentaddress: RD 2, Box219, Louisa,VA 23093. great deal of investigation using gel electropho- 14441 SCHIFFORNZS GROUP PHYLOGENY 445 resis(Zink and Johnson 1984, S. M. Lanyon 1985, tory traits is discussedin light of this new phy- Lanyon and Lanyon 1986, Johnson and Marten logenetic hypothesis. 1988), DNA-DNA hybridization (Sibley and Ahlquist 1985, Sibley et al. 1985) and cladistic REVIEW OF SYSTEMATIC HISTORY OF analysis of a variety of morphological characters SCHZFFORNZS GROUP GENERA (Lanyon 1984a, 1984b, 1985, 1986, 1988a, The genusSchzjbrnis contains three specieswhich 1988b, 1988~; McKitrick 1985; Lanyon and range from southern Mexico to southeasternBra- Lanyon 1986; Prum, unpubl.). Of particular in- zil. The best known species,Schijbrnis turdinus, terest to this investigation, McKitrick (1985) is commonly known as the Thrush-like Man- identified a problematic clade of tyrannoids (in- akin. The genus was placed in the Pipridae by cluding SchiJbrnis, Pachyramphus, and Tityra) Sclater (1888) under the names Schzjix-nisand which shares derived morphological characters Heteropelma, based on the possessionof exap- both with cotingids and piprids and with tyrant sidean tarsal scutellation and extensive fusion of flycatchers.In an investigation of the monophyly the outer toes (digits III and IV) which he used of the Pipridae and Cotingidae, Prum (unpubl.) to define the family. Sclater ( 188 8) placed Schif has expanded this group to include a number of fornis and several other genera, including Lani- other tyrannoid genera. isoma (seebelow), in the separatesubfamily Pti- We present here a cladistic analysis using mor- lochlorinae in recognitionof their differencesfrom phological charactersof the phylogeny of a pre- other typical piprids, such as dull, sexually viously unrecognized monophyletic assemblage monomorphic plumage and prominently hooked, of six genera in this problematic tyrannoid as- tyrannid-like bill. Ridgway (1907) and Hellmayr semblage. These genera are presently placed in (1929) left Schijbrnis in the Pipridae but did not three different tyrannoid families: Schzxornis recognizethe subfamilies of Sclater. Despite sug- (Pipridae), Laniisoma, Zodopleura(Cotingidae), gestionsthat Schzfirnis might belong in the Ty- Laniocera, Xenopsaris,and Pachyramphus(Ty- rannidae (Ames 197 1) or the Cotingidae (Meyer rannidae). The systematic relationships of these de Schauensee 1966, Wetmore 1972), Snow six genera have remained problematic or enig- (1975, 1979) followed previous authors and kept matic for over 100 years, resulting in frequent the genus in the Pipridae, citing the lack of sub- reallocation to different families or placement in stantive evidence supporting a specifictaxonom- isolated positions within them. Reevaluation of ic change. McKitrick (198 5) identified Schzjbr- their relationships has been made possible by nis as a member of a problematic group of genera advances in the understanding of the morpho- which possessboth a derived form of the femoral logical diversification of the tyrannoids (Warter artery sharedby cotingasand manakins, and de- 1965; Ames 1971; Lanyon 1984a, 1984b, 1985, rived internal syringeal cartilageswhich are pres- 1986, 1988a, 1988b, 1988~; McKitrick 1985; ent elsewhere only in tyrannids. S. M. Lanyon Prum, unpubl.), and by the recent collection of (1985) presented allozyme evidence for the the first anatomical and skeletal specimensof the placement of Schijbrnis as the sister group to genera Laniisoma and Xenopsaris. the piprids including Neopelma and Tyran- Following a summary of the taxonomic history neutes.Based on DNA-DNA hybridization, Sib- of these genera, which we refer to collectively as ley and Ahlquist (1985) placed Schiffornis, the Schifbrnis group, we present the morpho- Pachyramphus,and Tityra in the subfamily Ti- logical characterswhich support their monophy- tyrinae as the sister group to a restricted Tyran- ly and intergeneric relationships. We then discuss ninae. The remainder of the tyrannids were placed the diagnoses of the Schijbrnis group and its in the Mionectidae as the sister group to the rest monophyletic subgroups,the alternative hypoth- of the Tyrannoidea. Prum (unpubl.) recom- esesfor the relationships of the Schljbrnis group mended the removal of Sch$%ornisand five other genera, and the position of the Schzfirnis group genera of piprid-like tyrannoids (Neopelma, Ty- within the Tyrannoidea. We compare this mor- ranneutes,Neopipo, Piprites, and Sapayoa) from phological hypothesisto previous phylogenies of the Pipridae based on a cladistic analysis of mor- the tyrannoids based on DNA-DNA hybridiza- phological characters.All six of thesegenera lack tion (Sibley and Ahlquist 1985) and allozyme the derived, dorsal fusion of the Bl and B2 sy- electrophoresis(S. M. Lanyon 1985). Lastly, the ringeal elements which is shared by all other tra- evolution of various behavioral and natural his- ditional piprid genera (Prum, unpubl.). 446 RICHARD 0. PRUM AND WESLEY E. LANYON Laniisoma elegans, commonly called the cards, which range throughout the Neotropics to Shrike-like Cotinga, is the only species in this the southern United States. Pachyramphus and little-known genus. It has a disjunct distribution Platypsaris (originally a separate genus for the in southeasternBrazil and the eastern slopes of aglaiae species group) have been consistently the Andes from Bolivia to Venezuela (Snow placed in the Cotingidae near Tityra, on the basis 1982). Sclater (1888) placed Laniisoma (=Ptilo- of their possessionof emarginate or truncate 9th chloris) in the piprid subfamily Ptilochlorinae primaries (Sclater 1888, Ridgway 1907, Hell- next to Schzjknis, but Ridgway (1907) subse- mayr 1929, Snow 1973). Snow(1973) considered quently moved it to the Cotingidae, despite its the aglaiae speciesgroup to be insufficiently dis- exaspidean tarsi and