Ibis (2001) 143,307-309

Short communication of most and manaluns, in which the main arteri- al supply to the hindlimb is via the femoral artery (A. femoralis). In contrast, the main hindlimb arteries of both species of Andean green pihas, Lipaugus subalaris and L. A new genus for the Andean cryptolophus, were ischiadic (A. ischiadica), an apparent reversal within cotingas to the primitive state found in Green Pihas (Cotingidae) most that also occurs in the genera Rupicola, RICHARD 0. PRUM Phoenicircus, Carpornis, Pipreola, Ampelioides and Department of Ecology and Evolutionary Biologx and Natural Oxyrums (Prum 1990). Subsequent research on cotinga History Museum, and Dyche Hall, University of Kansas, syringeal morphology (Prum unpubl. data) has revealed Lawrence, KS 66045-2454, USA additional phylogenetically informative morphological differences between the Andean green pihas and the other members of Lipaugus. Most recently, a phylogenetic analysis of DNA sequences from a mitochondria1 protein The diverse Neotropical suboscine account for coding gene, cytochrome b (Prum et al. ZOOO), further more than a tenth of all avian species. Most radiations supports the conclusion that the genus Lipaugus as tradi- within this group have received little systematic scrutiny tionally constituted (Hellmayr 1929, Snow 1979) is since the historically standard classifications of Hellmayr polyphyletic. (Cory & Hellmayr 1924, 1925, 1927, Hellmayr 1929). In Prum (19903, I suggested that the species subalaris Recent advances in phylogenetic analysis and molecular and qptolophus be removed from Lipaugus Boie 1828, systematics have produced exciting new opportunities for but I placed them erroneously in the genus Lathria understanding higher-level phylogenetic relationships, Swainson 1837, which had been applied to these piha species limits and evolutionary history of the Neotropical species in some previous classifications (e.g. Sclater suboscine clade. 1888). Subsequently, D. F. Stotz (in litt.) brought to my To maintain congruence between avian classification attention that the type Lathria Swainson 1837 is and developments in phylogenetic research, it is impor- Muscicapa plumbea Lichtenstein = Muscicapa vociferans tant to revise avian to reflect new, well Wied which is also the type species of the genus Lipaugus. supported systematic findings. Often a traditional classifi- Stotz correctly observed that the genus Lathria is thus not cation can be made phylogenetically meaningful with a available for subalaris and cryptolophus if the type of minimum of changes: e.g. synonymizing a monotypic Lathria remains in Lipaugus. He further opined that genus that renders another genus paraphyletic or ahistor- another junior synonym of Lipaugus, Turdampelis Lesson ical. Some results, however, require that new names be 1844, was unavailable for a genus including subalaris and created to classify previously unrecognized clades accu- cryptolophus alone, because the type of Turdampelis is rately. Turdampelis lanioides Lesson, which I suggest should be The cotingas (Cotingidae) are a diverse Neotropical included within Lipaugus. Thus, no genus name is avail- family. Although the limits of the family have been in dis- able to include the newly recognized clade of cryptolophus pute for years, they are currently recognized to include and subalaris, and there are no strong data to support the traditional genera of Snow (1979), the plantcutters their placement in any other available cotingid genus. (Phytotoma) (Lanyon & Lanyon 1988, Prum et al. 2000), Herein, I propose a new genus name to include the Sharpbill (Oxyruncus) (Sibley et al. 1985, Sibley & species of this distinct clade of Andean green pihas: Ahlquist 1990, Prum et al. ZOOO), the tityras (Tityra) (Prum 1990, Prum et al. 2000) and the Schiffomis group gen. nov. of I’rum and Lanyon (Prum & Lanyon 1989, Prum et al. 2000) , , , Pachyramphus, Type species. Lipaugus subalaris Sclater. 1861. Proceedings Xenopsaris and lodopleura. of the Zoological Society of London, p. 210. Ecuador, Rio Snow (1982: 11 1) was first to suggest that the Andean Napo. green pihas - subalaris and cryptolophus - may be distant- Diagnosis. The genus is diagnosed by a combination of ly related to the rest of the piha species within the traits: (1) the insertion of the M. tracheolateralis on the traditional genus Lipaugus - vociferans, unirufus, fus- lateral syringeal membranes between the A1 and B1 cocinereus, lanioides, and streptophorus. Subsequently, in a syringeal supporting elements, (2) the absence of intrinsic test of the monophyly of the cotingas, Prum (1990) doc- syringeal musculature, (3) mainly ischiadic hindlimb arte- umented that Lipaugus species were morphologically rial supply, (4) largely olive-green plumage, with yellow quite divergent. The hindlimb arteries of Lzpaugus vocifer- eye ring, and yellow underwing coverts, and (5) partially ans revealed the derived heteromerous condition, typical concealed black crown patch in male or both sexes. Etymology. The genus Snowornis is named in honour of Dr David W. Snow for his many contributions to our Ernail: [email protected] understanding of the ecology, behaviour, and systematics

@ 2001 British Ornithologists’ Union 308 R.O. Prum

of the cotingas and manaluns, and for his original insights numerous other features from those of the members of into the role of frugivory in the evolution of polygyny and Lipaugus (Prum pers. obs.). Both species of Snowomis are iek behaviour. much less vocal than Lipaugus species. The vocali7ation of S. subalaris has been described as a 'clear and ringing two- DISCUSSION noted whistle: churrrrrrr-ee!' (Ridgely & Tudor 1994). The vocalizations of S. cyptolophus are unknown. The diagnosis of the new genus Snowornis includes the A recent phylogenetic analysis of sequences from the species S. subalaris and S. clyptolophus, and excludes all mitochondrial protein coding gene cytochrome b support- other cotinga species. The recognition of Snowomis is jus- ed a single phylogeny for 32 species representing 26 tified by data from arterial and syringeal morphology, genera of cotingas (Fig. 1) (Prum et al. 2000). The phy- plumage, and mitochondrial DNA sequences that all sup- logeny includes four main clades of cotingas which may port the monophyly of Snowomis and its distinctness from be recognized as subfamilies: Tityrinae, Phytotominae. L ipaugus. Rupicolinae and Cotinginae. The analysis included four The members of Snowomis are easily diagnosed by the species of pihas representing both genera: S. subalaris, 5. presence of largely olive-green plumage, yellow eye ring, cyptolophus, L. unirufus and L. fuscocinereus (Fig. 1). yellow underwing coverts and a partially concealed black Among the 3 2 cotinga species analysed, the percentage crown (Snow 1982). This black crown is found in both sequence divergence over the 375 base pairs from the 3' sexes of adult cyptolophus, but only in male subalaris. end of the mitochondrial cytochrome b gene analysed var- None of these plumage features occurs in Lipaugus. The crown patch is clearly derived within cotingas and consti- Ampefron rubrocnstafus Dohornrs scfaten tutes a distinct synapomorphy of the new genus. Rupicofa peruvana Snowomis subalaris can be further distinguished by its Rupicofa ruprcofa grey tail, undertail coverts and belly, whereas S. cyptolo- Phoenrcrrcus nigricoliis Pipieofa aicuafa phus is characterized by entirely green tail and underparts. Rupicolinae Pipreofa chloiolepidofii Since Prum (1990), 1 have made additional observa- oxyruncus cr,sta1us tions of the hindlimb arterial supply in Lipaugus. In Arnpefrordes fschudri addition to L. vociferans, the derived femoral artery con- Carpodectes hopke, dition is also present in unirufus, fuscocinereus and XJphoiena punicea Gymnoderus foelidus - srreptophorus. (As yet, no spirit specimens are available for Snowornis cryptolophus L. lanioides or Chiroqlla uropygialis, Wood et al. 1982.) - Snowornis subalaris So, the derived femoral artery is uniformly present in all Poiphyrolaema porphyiolaerna available members of Lipaugus and absent in Snowomis. Conroptifon mcrfhennyr Gephafopterus ornafus The absence of the femoral state in Snowomis is a derived Penssocephafus frrcoior loss that is probably synapomorphy of Snowomis or a Pyrodeius scutatus - synapomorphy shared by Snowomis (Prum et al. 2000). Oue,uia piiipurata Recent syringeal observations document that Haemafoderus miirtarrs - Lipaogus unirufus Cotinginae Lipaugus vociferans, L. unirufus, L. fuscocinereus, and L. Lipaogus fuscocinereus streptophorus all have a pair of intrinsic syringeal muscles cotinga cayana that originate on the ventral surface of the syringeal ele- Procnias afba ment A4 and insert on the Al/Rl lateral membranes lodopleura isabellae Laniisoma elegans (l'rum pers. obs.). These intrinsic muscles are derived Pachyiamphus marginatus xj-ithin cotingas and a synapomorphy of Lipaugus. The Pachyiamphus versicofoi loud and highly modulated vocalizations of L. vociferans Schrffoinrs major (the ), L. unirufus, L. fuscocinereus and Tityia mquisrlor rityia cayana possibly L. streptophorus (Kidgely & Tudor 1994) imply that these intrinsic muscles function directly in vocal P,ppntes chioris niodulations. Similar vocalizations \vith rapid, intense vol- Pipiids (5 Species1 ume modulations are known from L. lanioides and Chirocylla uropygialis (Ridgely & Tudor 1994), for which no syringcal specimens are available. Rased on their vocal Figure 1. Single most parsimonious phylogenetic hypothesis for behaviour, however, it appears likely that L. lanioides and 32 cotinga species based on 375 base-pair sequences from the C uropygiialis also have these derived intrinsic muscles 3' end of the mitochondria1 protein coding gene cytochrome b (Prum eta/. 2000). This molecular phylogenetic hypothesis sup- and are members of the monophyletic Lipaugus clade. In ports the monophyly of Snowornis, the monophyly of Lipaugus contrast, the syringes of Snowomis subalaris and S. cyp- excluding S. subalaris and S. cryptolophus, and a distant phylo- /olophu.s lack intrinsic syringeal muscles, which is the genetic relationship between Snowornis and Lipaugus within the general plesiomorphic condition for cotingas. The syringes subfamily Cotinginae. Species of Snowornis and Lipaugus are of S. subalaris and S. cyptolophus are also quite distinct in shown in boldface.

2001 British Ornithologists' Union, Ibis, 143, 307-309 New Genus for Andean Green Piha 309

ied from 4.3%(Cephalopterus omatus vs. Perissocephalus This paper is dedicated to Dr David W. Snow for his inspirational work on the display behaviour, breeding systems and evolution of tricolor) to 25.7% (Carpodectes hopkei vs. Laniisoma ele- manaluns and cotingas. Douglas F. Stotz kindly pointed out the gans). Percentage observed sequence divergence between taxonomic error in my earlier paper (Prum 1990). Critical mole- S. subalaris and S. cryptolophus was 10.8O/0 and between L. cular and morphological specimens for this work were provided unirufus and L. fuscocinereus was 12.8%. In contrast, the by the Academy of Natural Sciences of Philadelphia, American average percentage sequence divergence between the Museum of Natural History, British Museum (Natural History), species of Lipaugus and Snowomis was 20.2% Field Museum of Natural History, Louisiana State University (19.0-2 1.2%). Furthermore, the most parsimonious phy- Museum of Natural Science, and US National Museum of logenetic hypothesis for these data identified both Natural History. The manuscript has been improved by com- ments from K. Bostwick, S. Hilty, A.T. Peterson, M. Robbins, K. Lipaugus and Snowomis as exclusive monophyletic Zyskowski, and two anonymous reviewers. This research was groups, and both of these clades were more closely relat- supported by a grant from the National Science Foundation ed to other lineages within the subfamily Cotinginae (Fig. (DEB-93 18273). 1). Many of the specific branches within the Cotinginae in this hypothesis of phylogeny are not strongly supported (Prum et al. 2000). For example, there is little support for REFERENCES a sister group relationship between Snowornis and Poqhyrolaema. However, the molecular data do not sup- Cory, C.B. & Hellmayr, C.E. 1924. Catalogue of birds of the Amer- port any close relationship between Snowomis and icas, part 3. Pteroptochidae - Conopophagidae - Formicariidae. Lipaugus. Pub/. Field Mus. Nat. Hist. 223: 1-369. In summary, derived features of the syrinx and molecu- Cory, C.B. & Hellmayr, C.E. 1925. Catalogue of birds of the Amer- icas, part 4. Furnariidae-Dendrocolaptidae. Pub/. Field Mus. lar sequences support the monophyly of Lipaugus Nat. Hist. 234: 1-390. excluding subalaris and cryptolophus. As suggested by Cory, C.B. & Hellmayr, C.E. 1927. Catalogue of birds of the Amer- Remsen et al. (1982), Chirocylla uropygialis is distin- icas, part 5. Tyrannidae. Publ. Field Mus. Nat. Hist. 242: 1-517. guished only by its autapomorphous wing feather Hellmayr, C.E. 1929. Catalogue of birds of the Americas, part 6. specializations, and should be placed within Lipaugus. Oxyruncidae-Pipridae-Cotingidae-Rupicolidae-Phytotomidae. Lathria, Turdampelis, and Chirocylla are all junior syn- Publ. Field Mus. Nat. Hist. 266: 1-258. onyms of Lipaugus. Derived plumage traits and molecular Lanyon, S.M. & Lanyon, W.E. 1988. The systematic position of the sequences support the monophyly of Snowomis, including Plantcutters, Phytotoma. Auk 106: 422-432. only subalaris and cryptolophus. Additional variation in Prum, R.O. 1990. A test of the monophyly of the manakins (Pipri- hindlimb arteries and molecular sequences further sup- dae) and of the cotingas (Cotingidae) based on morphology. Occ. Pap. Mus. Zool. Univ. Michigan 723: 1-44. ports the conclusion that these two lineages of pihas are Prum, R.O. & Lanyon, W.E. 1989. Monophyly and phylogeny of the not closely related within the Cotinginae. Although the Schiffornis group (Tyrannoidea). Condor 91 : 444-4131, current hypotheses for the relationships among Lipaugus, Prum, R.O., Rice, N.H., Mobley, J.A. & Dimmick, W.W. 2000. A Snowomis, and other cotinga genera are by no means final, preliminary phylogenetic hypothesis for the cotingas (Cotingidae) data from different sources corroborate that the two gen- based on mitochondria1 DNA. Auk 117: 236-241. era of pihas are not closely related. Remsen, J.V. Jr, Parker, T.A., 111 & Ridgely, R.S. 1982. Natural his- Snowomis subalaris and S. cryptolophus remain poorly tory notes on some poorly known Bolivian birds. Gerfaut 72: known in nature (Snow 1982, Ridgely & Tudor 1994). 77-87. The few reported observations indicate that these species Ridgely, R.S. & Tudor, G. 1994. The Birds of South America, Vol. 2. are generally quiet denizens of humid upper tropical and The Suboscine Passerines. Austin, TX: University of Texas Press. humid lower montane forests of the Andes where they Sibley, C.G. & Ahlquist, J.E. 1990. Phylogeny and classification of feed on both insects and fruit. S. subalaris apparently pro- birds. New Haven: Yale University Press. duces a clear two-note whistled call (Snow 1982, Ridgely Sibley, C.G., Lanyon, S.M. & Ahlquist, J.E. 1985. The relation- & Tudor 19943, but there are no indications of lek behav- ships of the sharpbill (Oxyruncus cristatus). Condor86: 48-52. iour in either species, as in many Lipaugus (Snow 1982, Snow, D.W. 1979. Tityrinae, Pipridae, Cotingidae. In Traylor, M.A. Jr Ridgely & Tudor 1994). However, Snowomis belongs in (ed.) Check-list of Birds of the World, a Continuation of the Work Cotinginae which includes many other polygynous of James L. Peter, Vol. 8: 229-308. Cambridge, MA: Museum of species. Further research on the natural history and phy- Comparative Zoology. logenetic relationship of the Andean green pihas will Snow, D.W. 1982. The Cotingas. Ithaca, NY: Cornell University further elucidate the patterns of morphological, behav- Press. Wood, D.S., Zusi, R.L., & Jenkinson, M.A. 1982. World lnventory ioural and ecological evolution within the cotingas. Lastly, of Avian spirit specimens. Norman, OK: American Ornithologists' continuing to refer to S. subalaris and S. cryptolophus as Union. pihas would merely perpetuate the misimpression that these species are closely related to Lipaugus. Therefore, I recommend that S. subalaris and S. cryptolophus be referred to in English as the Grey-tailed Cotinga and the Olivaceous Cotinga respectively. Received 8 September 1999; revision accepted 19 June 2000

@ 2001 British Ornithologists' Union, Ibis, 143, 307-309