TITYRAS, MOURNERS & ALLIES
- 30 species in the Neotropics
- DR personal total: 13 species (43%), 1
uncertainty has long existed about the relationship of the suboscine
passerines in the Neotropics. There were two major groups — the
Tyrannida and the Furnariida (e.g., Sibley & Ahlquist 1990) — but
the relationships of the families within these two groups were
problematic. The most difficult was trying to find the boundaries
between Tyrant Flycatchers, Manakins, and Cotingas. Writing in his
introduction to a book on Cotingas, Snow (2004) wrote: "Following a
number of nineteenth-century anatomical studies, these three families
were distinguished from one another on the basis of the shape of the
bill and the scutellation of the tarsus, as well as whether or not the
toes were untied and, if they were, to what extent. It soon became
clear, however, that a classification based on these characters
contained awkward exceptions."
studies have sorted out a good number of this "awkward exceptions." It
turns out there are not just three families within the Tyrannida, but
four! The fourth family contains at least 8 genera for which convergent
evolution had masked their origins. Among them are four species of
tityras, including Masked Tityra (left or above; in a
nice shot by Greg Lasley that just happens, incidentally, to document
the first U.S. record of the species!)
on a congruence of multiple studies of molecular and morphological
characters, Ericson et al. (2006) formally proposed that the new family
be called the Tityridae. Included are species long assigned to Tyrant
Flycatchers (e.g., the tityras, to Cotingas (e.g., purpletufts), and to
Manakins (e.g., the genus Schiffornis).
There still remain some "difficult" species that have not yet been
sorted out, but the main outlines of family Tityridae is now reasonably clear.
* An assemblage of 23 species of mourners (genus Laniocera), becards (Pachyramphus), and tityras (Tityra).
Over a decade ago, Sick (1993) wrote: "Their syringes and other
characters, as well as their ecology, indicate closer relationship to
the tyrannids than to the cotingids. For me, knowing these birds in
life, it was a relief to remove them from the cotingids, especially
considering that their nest types do not occur among cotingids." Most
previous authorities had placed this group with tyrant flycatchers;
* The 3 species Schiffornis mourners, the 3 species of purpletufts (Iodopleura) and two species in monotypic genera: White-naped Xenopsaris Xenopsaris albinucha and "Shrike-like Cotinga," now usually called Elegant Mourner (Laniisoma elegans). Together these, sometimes along with the 3 species of piprites (Piprites),
have been called the "Schiffornis assemblage." Biochemically they form
a monophyletic group (e.g. Lanyon 1985, Prum & Lanyon 1989, Prum et
al. 2000, Chesser 2004) but there is now doubt that the piprites are
included (see Ericson et al. 2006; Piprites may by tyrant flycatchers); and
* One to three odd, monotypic genera: Kinglet Calyptura (Calyptura cristata; once thought extinct but recently rediscovered and then it disappeared again!), the enigmatic Sharpbill (Oxyruncus cristatus; more below), and Swallow-tailed Cotinga (Phibalura flavirostris).
As to these final three unique species there is insufficient evidence
to firmly place them anywhere. Sharpbill was proposed to be within this
family by Ericson et al. (2006) and it is clearly not a cotinga (Ohlson
et al. 2007), but some doubts remain. I follow Ericson et al. (2006),
though, in assigning it to this family. The Kinglet Calyptura and
Swallow-tailed Cotinga have just not been studied enough, so I continue
to assigned Calyptura and Phibalura to their traditional family, the Cotingidae.
information will be developed over time. It will be particularly
interesting to see where the Kinglet Calyptura and Swallow-tailed
Cotinga fit in. No one has published any molecular evidence about Calyptura — it remains essentially a mystery. Dickinson (2003) put Phibalura
near the others in this family, but this may be based on unpublished
evidence. For now, because I have to put them somewhere, I retain them
most members of the Tityridae are primarily fruit-eaters, others seem
to be primarily insectivores. Although DNA evidence shows that this is
a distinctive evolutionary grouping, many have evolved to fit specific
niches. The purpletufts, for example, live in small parties at the
forest edge (as along rivers) and eat fruit, like mistletoe. In
contrast, many becards sally after insects in the subcanopy. One
example is Rose-throated Becard (left), a species
that ranges from the southwestern United States to Costa Rica. This
photo is from the latter country, where populations do not have rose on
Tityras can be conspicuous and
aggressive. Rather little is known about a number of the mourners, or
the Xenopsaris of the Amazon basin. There is still a lot to be learned
about this 'new' family.
Photos: Greg Lasley photographed the Masked Tityra Tityra semifasciata
at Bentsen-Rio Grande S.P., Texas, on February 20, 1990, where it
represented a first U.S. record [various shots in Greg's photo series
of this bird have been published in American Birds 44: 223 (1990) and Birding 24: 280 (1992)]. The male Rose-throated Becard
Pachyramphus aglaiae was photographed at Rio Tigre, Osa Peninsula,
Costa Rica, on 26 Dec 2007. Photo of becard
© Don Roberson; photo of tityra © 2007
Greg Lasley, and are used with
all rights reserved.
Bibliographic note: There is no "family book"
per se but members of the family have been covered in Handbook of the Birds of the World, but under various previously-assigned families. Vol. 9 of HBW
placed the 23 becards/tityras with Tyrant Flycatchers (Tyrannidae),
puts the Schiffornis assemblage in the Manakins (Pipridae), and
sequenced the Purpletufts, and Elegant Mourner, Kinglet Calyptura,
Sharpbill, and Swallow-tailed Cotinga in the Cotingas (e.g., Snow 2004).
- Chesser, R.T. 2004. Molecular systematics of New World suboscine birds. Molecular Phylogenetics and Evolution 32: 11-24.
Dickinson, E.C., ed. 2003. The Howard & Moore Complete Checklist of
the Birds of the World. 3d ed. Princeton Univ. Press, Princeton, N.J.
P.G.P., D. Zuccon, J.I. Ohlson, U.S. Johansson, H. Alvarenga, and R.O.
Prum. 2006. Higher-level phylogeny and morphological evolution of
tyrant flycatchers, cotingas, manakins, and their allies (Aves:
Tyrannida). Molec. Phylogenetics Evol. 40: 471-483.
Johansson, U.S., T.J. Parsons, M. Irestedt, and P.G.P. Ericson. 2001.
Clades within "higher land birds," evaluated by nuclear DNA sequences.
J. Zool. Syst. Evol. Research 39: 37-51
- Lanyon, S.M. 1985. Molecular perspective on higher-level relationships in the Tyrannidae (Aves). Syst. Zool. 34: 404-418.
Lanyon, W.E. 1967. Revision and probable evolution of the Myiarchus
flycatcher of the West Indes. Bull. Amer. Mus. Nat. Hist. 136: 329-370.
J.I., R.O. Prum, and P.G.P. Ericson. 2007. A molecular phylogeny of the
cotingas (Aves: Cotingidae). Molec. Phylogenetics Evol. 42: 25-37.
- Prum, R.O., and W.E. Lanyon. 1989. Monophyly and phylogeny of the Schiffornis group (Tyrannoidea). Condor 91: 444-461.
Prum, R.O., N.H. Rice, J.A. Mobley, and W,W. Dimmick. 2000. A
preliminary phylogenetic hypothesis for the cotingas (Cotingidae) based
on mitochondrial DNA . Auk 117: 236-241.
- Ridgely, R.S., and G. Tudor. 1994. The Birds of South America. Vol. 2: The Suboscine Passerines. Univ of Texas, Austin.
Snow, D.B.. 2004. Family Cotingidae (Cotingas), pp. 32-108 in Del Hoyo,
J. Elliott, A., & Sargatal, J. eds. Handbook of the Birds of the
World. Vol. 9. Lynx Edicions, Barcelona.
- Sibley, C.G., and J.E. Ahlquist. 1990. Phylogeny and Classification of Birds of the World. Yale Univ. Press, New Haven, CT.
Sick, H. 1993. Birds in Brazil: A Natural History. Translated from
Portuguese by W. Belton. Princeton Univ. Press, Princeton, N.J.
- South American Checklist Committee. 2007. On-line; Remsen et al,, eds. A subcommittee of the American Ornithologists' Union.
Page created as part of Cotingidae on 8-9 Dec 2004, new family page created 4-5 Nov 2007, slightly revised 27 Aug 2011