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SHRIKE-BABBLERS Pteruthiidae |
- 9 species in south Asia
- DR personal total: 3 species (33%), 1 photo'd
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The
Shrike-babblers are a small group of south Asian birds, traditionally
considered to be "Babblers" in the old, traditional family Timaliidae.
One example is Green Shrike-babbler (left).
The old traditional "babbler" family is now split up into five new families [see the new page on the Timaliidae
for details], plus offshoot families of birds that have been proven,
though biochemical research, to be unrelated to babblers; Alström
et al. (2006), Cibois (2003, 2010).
Like another "babbler" once known as "White-bellied Yuhina Y. zantholeuca," both proved to be most closely related to New World vireos (Cibois 2003, Alström
et al. 2006, Reddy 2008). Truly shocking! Now called Erpornis Erpornis zantholeuca, it and Shrike-babblers are now often placed among the Vireonidae. |
The shrike-vireos (genus Pteruthius)
are indeed related to vireos (Reddy & Cracraft 2007, Reddy 2008).
They do have a unique bill (right; Green Shrike-Babbler again; all
photos on this page). Reddy & Cracraft's analysis, using two
species of shrike-vireos, sequenced two nuclear genes. They found that
shrike-vireos were the sister to a clade that included the North
American vireos and Erpornis. Reddy & Cracraft (2007), followed up
by Reddy (2008), calculated the divergence time of the shrike-babblers
from the remaining Vireonidae at 29.5 million years ago (plus or minus
2.5 million years). We can think of it as about 30 million years ago.
They also estimated the divergence of Erpornis from the New World
vireos at 23.8 million years ago (plus or minus 2.8 mya).
From
the standpoint of Family rank, one can view this evidence in three
ways: (1) lump all of them (vireos, Erpornis, shrike-babblers) into one
family (Vireonidae); (2) lump Erpornis with vireos and split
shrike-babblers as a separate family (the Pteruthiidae), or (3) place
each of the two Asian groups, both of them early offshoots of the
vireonid line, into separate families (Pteruthiidae, Erpornidae) and
leave the New World vireos in their own family (Vireonidae). Any of
these three approaches is justified by the current evidence. The first
approach emphasizes the relationships between these groups. The second
approaches emphasizes the long time of divergence of the
shrike-babblers. The third approach is a middle ground (chosen here)
emphasizing the offshoots that have been separate from vireos for ~24
million years. |
For
the moment, I chose the latter approach. It seems to me to be the most
consistent with the current approach to Family level taxa. For example,
Beresford et al. (2005) wrote, in reviewing the evolution of a group of
African warblers, that the "basal divergences among Old World warblers
range from 32.3 mya between the Spenoeacus clade and remaining warblers to 31.3 mya between megalurine and other warblers." The Spenoeacus clade mentioned is now widely considered a separate family, the Macrosphenidae [African warblers & crombecs], and the rest of the Old World warblers have been formally broken up into separate families, including the Locustellidae [grassbirds].
If these evolutionary clades are considered Families, having split off
from their nearest relatives ~31 to 32 million years ago, it seems
logical to me that the shrike-babblers — diverging in about that same
time frame at ~30 mya — are better assigned the rank of Family, rather
than Subfamily.
So, for the moment, shrike-babblers
are flying at you from your screen as a new family. Yet note the short,
rather broad and unusual bill and the large feet. Many vireos have
these same characteristics. In central China, we found that Green
Shrike-babbler was found in mixed species flocks — which could include
babblers, scimitar-babblers, parrotbills, warblers, and woodpeckers —
but that Green Shrike-babbler was the "core" species in the flock. It
was the "leader" and the rest of the flock coalesced around it. Its
vocalizations were a way to follow the flock through thickets and
bamboo.
There is yet another interesting research
study on shrike-babblers. Traditionally, most authorities have listed
five species of shrike-babbler.
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These five are shown on Hilary Burn's plate in Vol. 12 of the Handbook of the Birds of the World
(right), where shrike-babblers still appear among the old Timaliidae
(Collar & Robson 2007). These are (keyed to numbers on the plate):
- Black-headed Shrike-babbler P. rufiventer (297)
- White-browed Shrike-babbler P. flaviscapis (298)
- Green Shrike-babbler P. xanthochlorus (299)
- Black-eared Shrike-babbler P. melanotis (300)
- Chestnut-fronted Shrike-babbler P. aenobarbus (301)
Reddy
(2008) obtained tissue samples of all five traditional species and most
of the 20+ named subspecies, and undertook molecular phylogenetic
analyses and a systematic review of their morphology and plumage
characters. You can see in the HBW plate that there is much diversity.
As just one example, Green Shrike-babbler (#299), as traditionally
constituted, there are four named subspecies, two of which lack an
eyering [these are Himalayan populations, illustrated by nominate xanthochlorus on the plate] and two of which have an obvious eyering [these are Chinese populations; pallidus
is illustrated]. It is also apparent from the painting that some
shrike-babblers have strong sexual dimorphism [in contrast, New World
vireos lack sexual dimorphism, so shrike-babblers differ in that way].
Reddy's
(2008) work was an effort to determine the number of species that such
a thorough review might uncover, and she chose to use the Phylogenetic
Species Concept (PSC) for her results. In short, she determined that
the traditional 5 species should be divided into 19 PSC species.
The
problem with the PSC is that any single unique trait, even one that
lacks any reproductive significance, can be used to "identify" a
species. Many evolutionary biologists deplore the PSC concept when
applied to definitions of species (see Remsen 2005 for a good
overview), and the 'over-splitting' that it engenders can have
significant negative impacts on conservation work.
Fortunately,
as to shrike-babblers, Rheindt & Eaton (2009), undertook a thorough
review of Reddy's (2008) work, added vocalizations to the analysis, and
reviewed molecular, morphological, and vocal evidence using the
Biological Species Concept (BSC), which is by far the primary species
concept adopted worldwide. They recommended that the 5 traditional
shrike-babblers be expanded to 9 species, and that one other split is
likely [i.e., between the eye-ringed and unringed Green
Shrike-Babblers] but that evidence from nuclear DNA genetics is needed.
I suspect that the 5 traditional shrike-babblers will become 10
shrike-babblers in due course, but that is still just about half of the
19 "species" proposed by Reddy (2008). It is an interesting topic —
stay tuned.
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Photos: The Green Shrike-babbler Pteruthius xanthochlorus was at Foping Nature Reserve, Shaanxi, China, on 12 Nov 2010. All
photos © Don Roberson; all rights reserved. |
Bibliographic notes
There
is no "family book" on the Shrike-babblers (indeed, I am about the only one assigning them Family rank) but the Handbook of the Birds of the World
account (Collar & Robson 2007) is excellent, with great
photos, although it is not quite up with the most current
taxonomic findings, most of which regarding shrike-babblers postdate its publication.
Literature cited:
Alström
P., P.G.P. Ericson, U. Olsson, and P. Sundberg. 2006. Phylogeny and
classification of the avian superfamily Sylvioidea. Molecular
Phylogenetics & Evolution 38: 381-397.
Barker,
F.K., A. Cibois, P. Schikler, J. Feinstein, and J. Cracraft. 2004.
Phylogeny and diversification of the largest avian radiation. Proc.
Nat. Acad. Sci. 101: 11040-11045.
Cibois, A. 2003. Mitochondrial DNA phylogeny
of babblers (Timaliidae). Auk 120: 35-54.
Cibois,
A., M. Gelang, and E. Pasquet. 2010. An overview of the babblers and
associated groups. Systematic Notes on Asian Birds 68: 1-5.
Clements, J.F. 1991. Birds of the World: A Check-List. 4th ed. Ibis Publishing, Vista, CA.
Collar, N.J., and C. Robson. 2007. "Babblers (Timaliidae)," pp. 70-291 in Handbook of the Birds of the World (del Hoyo, J., A. Elliott & D.A.
Christie, eds). Vol. 12. Lynx Edicions, Barcelona, Spain.
Gelang,
M., A. Cibois, E. Pasquet, U. Olsson, P. Alström, and P.G.P.
Ericson. 2009. Phylogeny of babblers (Aves, Passeriformes): major
lineages, family limits and classification. Zoologica Scripta 38:
225-236.
Jønsson,
K.A., and J. Fjeldså. 2006. A phylogenetic supertree of oscine
passerine birds. Zoologica Scripta 35: 149-186.
Reddy,
S. 2008. Systematics and biogeography of the shrike-babblers
(Pteruthius): Species limits, molecular phylogenetics, and
diversification patterns across southern Asia. Molec. Phylog. Evol. 47:
54-72.
Reddy, S., and J. Cracraft. 2007. Old World Shrike-babblers (Pteruthius) belong with New World Vireos (Vireonidae). Molec. Phylog. Evol. 44: 1352-1357.
Remsen, J.V., Jr. 2005. Pattern, process, and rigor meet classification. Auk 122: 403–413.
Rheindt,
F.E., and J.A. Eaton. 2009. Species limits in Pteruthius (Aves:
Corvida) shrike-babblers: a comparison between the Biological and
Phylogenetic Species Concepts. Zootaxa 2301: 29–54.
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