New World Suboscines: Tyrannides
We turn to the New World suboscines and find that the avian tree is again divided into two parts: Tyrannida and Furnariida. As Tyrannida is the smaller group, it goes first.
New World Suboscines II: Tyrannida
The Tyrannida are the other group of New World suboscines. They have been variously separated into three to five families, sometimes including the Phytotomidae (plantcutters) and Oxyruncidae (sharpbill). The taxonomic position of a number of the Tyrannida has been called into question, starting with McKitrick (1985).
Recent studies have generally found four main clades: Pipridae (manakins), Cotingidae (cotingas), Tityridae (tityras and becards), Tyrannidae (tyrant flycatchers). The last is sometimes divided. However, other than argeement that they are each other's closest relatives, there is no consensus on how they relate. There are 18 possible fully resolved trees. Some analyses do not fully resolve the tree, but of those that do, at least five different versions have appeared in the recent literature. The major contenders are shown below.
The Ericson et al. tree is based on many genes, but relatively few species. The other two are based on DNA from many species, but fewer genes. As for the genes themselves: Ericson et al. include RAG-1, G3PDH, c-myc, cyt-b, myoglobin, and ODC. Ohlson et al. consider G3PDH, myoglobin, and ODC (ODC alone with Ericson's species gives the same result), while Tello et al. look at RAG-1 and RAG-2. The TiF list now uses the Ericson et al. arrangement.
Prum and Lanyon (1989) used morphological evidence to argue that Schiffornis, previously considered manikins, were actually part of a clade related to the becards and tityras. This has subsequently been supported by genetic evidence. Some of the other birds in the clade had previously been considered Tyrannidae. They also called attention to some additional problematic genera, including Neopipo, Piprites, Neopelma, Tyranneutes and Sapayoa that seemed misplaced.
It was eventually realized that Sapayoa is an Old World Suboscine. Neopelma and Tyranneutes are now known to be manakins. We will take the position that Neopipo and Piprites are relatively basal members of the Tyrannidae, although matters are far from settled.
A considerable amount of genetic analysis has been done on the Tyrannida. Significant recent papers include Tello and Bates, 2007; Ohlson et al., 2008; Rheindt et al., 2008; Tello et al., 2009.
Pipridae: Manakins
12 genera, 51 species HBW-9
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In the past, several would-be manakin species have turned out to be something else (e.g., Sapayoa). It now seems there is a core group of manakins, which consists of the species listed below except for a clade consisting of Neopelma and Tyranneutes, (see Rêgo et al., 2007). Although it has been suggested that the Neopelma/Tyranneutes clade belongs elsewhere, several independent genetic analyses (Barber and Rice, 2007; Rheindt et al., 2008a; Ohlson et al., 2008; Tello et al., 2009) have concluded they are sister to the core manakins. The comprehensive studies of Ohlson et al. and Tello et al. give us confidence that all of the members of the Pipridae have now been correctly identified.
The order here is now based on a tree diagram due to Hackett that was used by Anciães and Peterson (2009). The diagram is entirely consistent with Ohlson et al. (2008), and broadly consistent with Rêgo et al. (2007) and Tello et al. (2009), which I previously relied on.
This has led to another round of generic changes. Several Neopelma have be reclassified Tyranneutes, I merged Corapipo into Masius, separated Chloropipo from Xenopipo (moving X. holochlora to Lepidothrix), returned Dixiphia Ceratopipra to Pipra, into which I merged Machaeropterus.
- Wied's Tyrant-Manakin, Neopelma aurifrons
- Serra do Mar Tyrant-Manakin, Neopelma chrysolophum
- Dwarf Tyrant-Manakin, Tyranneutes stolzmanni
- Tiny Tyrant-Manakin, Tyranneutes virescens
- Pale-bellied Tyrant-Manakin, Tyranneutes pallescens
- Saffron-crested Tyrant-Manakin, Tyranneutes chrysocephalus
- Sulphur-bellied Tyrant-Manakin, Tyranneutes sulphureiventer
- Pin-tailed Manakin, Ilicura militaris
- White-throated Manakin, Masius gutturalis
- Golden-winged Manakin, Masius chrysopterus
- White-ruffed Manakin, Masius alterus
- White-bibbed Manakin, Masius leucorrhous
- Araripe Manakin, Antilophia bokermanni
- Helmeted Manakin, Antilophia galeata
- Swallow-tailed Manakin / Blue Manakin, Chiroxiphia caudata
- Lance-tailed Manakin, Chiroxiphia lanceolata
- Long-tailed Manakin, Chiroxiphia linearis
- Blue-backed Manakin, Chiroxiphia pareola
- Yungas Manakin, Chiroxiphia boliviana
- Jet Manakin, Chloropipo unicolor
- Yellow-headed Manakin, Chloropipo flavicapilla
- Olive Manakin, Xenopipo uniformis
- Black Manakin, Xenopipo atronitens
- Green Manakin, Lepidothrix holochlora
- White-fronted Manakin, Lepidothrix serena
- Orange-bellied Manakin, Lepidothrix suavissima
- Blue-crowned Manakin, Lepidothrix coronata
- Blue-rumped Manakin, Lepidothrix isidorei
- Cerulean-capped Manakin, Lepidothrix coeruleocapilla
- Snow-capped Manakin, Lepidothrix nattereri
- Golden-crowned Manakin, Lepidothrix vilasboasi
- Opal-crowned Manakin, Lepidothrix iris
- Orange-crowned Manakin / Orange-crested Manakin, Heterocercus aurantiivertex
- Yellow-crowned Manakin / Yellow-crested Manakin, Heterocercus flavivertex
- Flame-crowned Manakin / Flame-crested Manakin, Heterocercus linteatus
- White-bearded Manakin, Manacus manacus
- White-collared Manakin, Manacus candei
- Orange-collared Manakin, Manacus aurantiacus
- Golden-collared Manakin, Manacus vitellinus
- Wire-tailed Manakin, Pipra filicauda
- Crimson-hooded Manakin, Pipra aureola
- Band-tailed Manakin, Pipra fasciicauda
- Fiery-capped Manakin, Pipra pyrocephala
- Club-winged Manakin, Pipra deliciosa
- Striped Manakin, Pipra regulus
- White-crowned Manakin, Pipra pipra
- Scarlet-horned Manakin, Pipra cornuta
- Golden-headed Manakin, Pipra erythrocephala
- Red-capped Manakin, Pipra mentalis
- Red-headed Manakin, Pipra rubrocapilla
- Round-tailed Manakin, Pipra chloromeros
Cotingidae: Cotingas
26 genera, 66 species HBW-9
The Kinglet Calyptura and Swallow-tailed Cotinga
are traditionally placed in the cotingas, and that tradition
is followed here. The Kinglet Calyptura was long believed extinct
when rediscovered in the mid 1990's. Its future remains doubtful.
Although sometimes made a separate family, the Phytotomidae (plantcutters) are
also included in the Cotingidae.
The overall structure of Cotingidae is based on Ohlson et al. (2007). They did not include Calyptura or Phibalura in their analysis, but all other genera except Carpornis were included. Carpornis was included in Tello et al. (2009). Several well-supported clades are notable. The first is the fruiteater clade. The second is the plantcutter clade. The red cotingas, including the cocks-of-the-rock, also form a well-supported clade, which may be sister to the plantcutter clade. These two were joined by the Carpornis berryeaters and Snowornis pihas in Tello et al.'s analysis. Ohlson et al. came to a completely different conclusion regarding Snowornis, placing it as sister to the core cotingas (the remaining group), which also form a well-supported clade. Due to the lack of consensus about Snowornis, I've placed it better the two clades that have a claim on it. This is not the first time Snowornis has been moved around. It was once considered part of Lipaugus.
From here on, the branching pattern is not as clear-cut and may need some adjustment in the future. Only the last four genera are securely placed. However, it seems clear enough that all of these genera belong together. It's just the arrangement that is dicey. Following Ohlson et al. (2007), the next branch contains the fruitcrows and umbrellabirds. The remaining cotingas split into the blue cotingas (Cotinga), the pihas, and the “canopy cotingas”, including the bellbirds, one of the genera that may need to be moved when more information becomes available.
- Kinglet Calyptura, Calyptura cristata
- Swallow-tailed Cotinga, Phibalura flavirostris
- Scaled Fruiteater, Ampelioides tschudii
- Green-and-black Fruiteater, Pipreola riefferii
- Band-tailed Fruiteater, Pipreola intermedia
- Barred Fruiteater, Pipreola arcuata
- Golden-breasted Fruiteater, Pipreola aureopectus
- Orange-breasted Fruiteater, Pipreola jucunda
- Black-chested Fruiteater, Pipreola lubomirskii
- Masked Fruiteater, Pipreola pulchra
- Scarlet-breasted Fruiteater, Pipreola frontalis
- Fiery-throated Fruiteater, Pipreola chlorolepidota
- Handsome Fruiteater, Pipreola formosa
- Red-banded Fruiteater, Pipreola whitelyi
- Hooded Berryeater, Carpornis cucullata
- Black-headed Berryeater, Carpornis melanocephala
- Guianan Red-Cotinga, Phoenicircus carnifex
- Black-necked Red-Cotinga, Phoenicircus nigricollis
- Guianan Cock-of-the-rock, Rupicola rupicola
- Andean Cock-of-the-rock, Rupicola peruvianus
- White-cheeked Cotinga, Zaratornis stresemanni
- Peruvian Plantcutter, Phytotoma raimondii
- White-tipped Plantcutter, Phytotoma rutila
- Rufous-tailed Plantcutter, Phytotoma rara
- Chestnut-bellied Cotinga, Doliornis remseni
- Bay-vented Cotinga, Doliornis sclateri
- Red-crested Cotinga, Ampelion rubrocristatus
- Chestnut-crested Cotinga, Ampelion rufaxilla
- Gray-tailed Piha, Snowornis subalaris
- Olivaceous Piha, Snowornis cryptolophus
- Crimson Fruitcrow, Haematoderus militaris
- Purple-throated Fruitcrow, Querula purpurata
- Red-ruffed Fruitcrow, Pyroderus scutatus
- Capuchinbird, Perissocephalus tricolor
- Bare-necked Umbrellabird, Cephalopterus glabricollis
- Amazonian Umbrellabird, Cephalopterus ornatus
- Long-wattled Umbrellabird, Cephalopterus penduliger
- Lovely Cotinga, Cotinga amabilis
- Turquoise Cotinga, Cotinga ridgwayi
- Blue Cotinga, Cotinga nattererii
- Plum-throated Cotinga, Cotinga maynana
- Purple-breasted Cotinga, Cotinga cotinga
- Banded Cotinga, Cotinga maculata
- Spangled Cotinga, Cotinga cayana
- Black-and-gold Cotinga, Tijuca atra
- Gray-winged Cotinga, Tijuca condita
- Chestnut-capped Piha, Lipaugus weberi
- Dusky Piha, Lipaugus fuscocinereus
- Scimitar-winged Piha, Lipaugus uropygialis
- Rufous Piha, Lipaugus unirufus
- Screaming Piha, Lipaugus vociferans
- Cinnamon-vented Piha, Lipaugus lanioides
- Rose-collared Piha, Lipaugus streptophorus
- Purple-throated Cotinga, Porphyrolaema porphyrolaema
- Three-wattled Bellbird, Procnias tricarunculatus
- White Bellbird, Procnias albus
- Bearded Bellbird, Procnias averano
- Bare-throated Bellbird, Procnias nudicollis
- Black-faced Cotinga, Conioptilon mcilhennyi
- Bare-necked Fruitcrow, Gymnoderus foetidus
- Pompadour Cotinga, Xipholena punicea
- White-tailed Cotinga, Xipholena lamellipennis
- White-winged Cotinga, Xipholena atropurpurea
- Black-tipped Cotinga, Carpodectes hopkei
- Yellow-billed Cotinga, Carpodectes antoniae
- Snowy Cotinga, Carpodectes nitidus
Tityridae: Tityras, Becards
11 genera, 37 species Not HBW Family
The taxonomically troubling Schiffornis group has now grown into a
family: the Tityridae. The original Schiffornis group, which includes the tityras and becards,
forms the core of the Tityridae. This gathers together several genera
previously classified as Tyrannidae, Cotingidae, and Pipridae (see Barber and
Rice, 2007; Ohlson et al., 2008; Tello et al., 2009). There is general consensus
on how the core tityrids are organized. The main disagreement concerns whether
Laniisoma and Laniocera are closest relatives, as in Ohlson et al.,
and Tello et al., or whether Laniocera is closer to Schiffornis, as
found by Barber and Rice. Within the family, Schiffornis,
Laniocera, and Laniisoma form one clade, which is sister to a
clade consisting of the rest of the Tityridae, including becards
(Pachyramphus) and tityras (Tityra).
However, it seems that these are not all of the Tityridae. Although not all analyses agree, many recent papers (e.g., Ericson et al., 2006; Fjeldså et al., 2003; Ohlson et al., 2007; 2008; Tello et al., 2009) have indicated that the Sharpbill is the sister taxon to the core Tityridae. Accordingly, the Sharpbill is included in the Tityridae rather than being placed in its own monotypic family.
Tello and Bates (2007) discovered a small clade, the “Myiobius assemblage”, hidden within the Tyrannidae. Both they and Rheindt et al. (2008a) found evidence that this clade was a early branch in the Tyrannida, but did not sample enough basal taxa to know where to put it. A new paper by Ohlson et al. (2008) samples more of the relevant taxa. Their results suggest that the “Myiobius assemblage” is a basal clade in the Tityridae. The evidence they found for this is not strong, but it is the best that is currently available. Tello et al. (2009) also include the Myiobius assemblage in the Tityridae, but found it sister to the Sharpbill, with the whole lot sister to the core Tityridae. Because of that, I have put the Sharpbill as a separate branch between the Myiobius assemblage and the core Tityridae.
I've followed AOU's NACC in treating Sulphur-rumped Flycatcher, Myiobius sulphureipygius, and Whiskered Myiobius, Myiobius barbatus as separate species. SACC lumps them into one. Both have recently decided based on the same evidence (or lack thereof). I don't know which is right. The race mastacalis may also deserve specific status, but again, the evidence is weak.
- Royal Flycatcher, Onychorhynchus coronatus
- Ruddy-tailed Flycatcher, Terenotriccus erythrurus
- Tawny-breasted Flycatcher / Tawny-breasted Myiobius, Myiobius villosus
- Sulphur-rumped Flycatcher / Sulphur-rumped Myiobius, Myiobius sulphureipygius
- Whiskered Flycatcher / Whiskered Myiobius, Myiobius barbatus
- Black-tailed Flycatcher / Black-tailed Myiobius, Myiobius atricaudus
- Sharpbill, Oxyruncus cristatus
- Shrike-like Cotinga / Elegant Laniisoma, Laniisoma elegans
- Speckled Mourner, Laniocera rufescens
- Cinereous Mourner, Laniocera hypopyrra
- Varzea Schiffornis, Schiffornis major
- Thrush-like Schiffornis, Schiffornis turdina
- Greenish Schiffornis, Schiffornis virescens
- White-browed Purpletuft, Iodopleura isabellae
- Dusky Purpletuft, Iodopleura fusca
- Buff-throated Purpletuft, Iodopleura pipra
- Masked Tityra, Tityra semifasciata
- Black-crowned Tityra, Tityra inquisitor
- Black-tailed Tityra, Tityra cayana
- White-naped Xenopsaris, Xenopsaris albinucha
- Glossy-backed Becard, Pachyramphus surinamus
- Cinnamon Becard, Pachyramphus cinnamomeus
- Chestnut-crowned Becard, Pachyramphus castaneus
- Black-capped Becard, Pachyramphus marginatus
- Gray-collared Becard, Pachyramphus major
- White-winged Becard, Pachyramphus polychopterus
- Black-and-white Becard, Pachyramphus albogriseus
- Barred Becard, Pachyramphus versicolor
- Green-backed Becard, Pachyramphus viridis
- Yellow-cheeked Becard, Pachyramphus xanthogenys
- Slaty Becard, Pachyramphus spodiurus
- Cinereous Becard, Pachyramphus rufus
- Crested Becard, Pachyramphus validus
- Pink-throated Becard, Pachyramphus minor
- One-colored Becard, Pachyramphus homochrous
- Rose-throated Becard, Pachyramphus aglaiae
- Jamaican Becard, Pachyramphus niger
