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BABBLERS Timaliidae
Laughingthrushes, Typical Babblers & White-eyes |
- 336 species in Old World tropics
- DR personal total: 213 species (63%), 18 photos
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The
true Babblers are a huge family of (mostly) tropical forest birds in
the Old World. There is much diversity within the family. A good chunk
(50 species; 19%) are called laughingthrushes in the genus Garrulax. These are rangy, mid-sized, floppy-tailed landbirds like Streaked Laughingthrush
(left) of the Himalayan mid-elevations. Streaked Laughingthrush is easy
to observe as it forages in pairs in open rocky areas in the mountains;
its foraging behavior, its size, and the fact one finds it mostly in
closely-knit pairs all resemble the California Towhees Pipilo crissalis
in my home county. Many other laughingthrushes, though, are elusive
species in thick jungle, best located by voice. On a multi-week trip to
India, for example, we heard noisy flocks of the striking and
widespread White-crested Laughingthrush Garrulax leucolophus
several times — sometimes quite close — and yet Rita and I only saw
them on our next-to-final day and then only in flight across a small
densely vegetated canyon. All the other times it sounded like they were
laughing at us! |
We
are only slowly learning how to define babblers. It has long been an
ornithological joke that the Timaliidae was the "dust-bin" for Old
World birds of uncertain origin: in 1896 Alfred Newman called babbler
taxonomy "a festering mass" (Collar & Robson 2007). Studies in the
mid-2000s began to clarify the situation (e.g., Cibois 2003,
Alström et al. 2006, Jønsson & Fjeldså 2006) but
there is uncertainty whether the huge babbler assemblage should be
allocated to one or two families. Genetic evidence revealed that the Sylvia
warblers — the original, base genus in the 'Old World Warblers'
Sylviidae — is more closely related to Babblers than other Old World
Warblers [see a discussion of the break-up of the Old World Warblers]. Genetic evidence also showed that white-eyes, such as this Silver-eye
(right, shown eating a crab-apple on Heron I. which is part of
Australia's Great Barrier Reef), were also babblers. Alström et
al. (2006) would apparently combine all the obvious babblers with the
Sylvia warblers, the white-eyes, the parrotbills, and the Wrentit of
the New World to form one huge Timaliidae Family. Under that approach,
the family would have about 390 species. Jønsson &
Fjeldså (2006) took a slightly different approach. They divide
this huge group into two "clades" [evolutionary lineages] that are each
others closest relatives. If one were to assign Family rank to each
"clade," there would be the Babbler Family [Timaliidae] of ~340
species, which include all the White-eyes once considered to be a
separate family, and a second Family that includes the Sylvia
'warblers', the Wrentit, various babblers, and almost all of the
parrotbills (also once considered a separate family). I like the latter
approach for the moment; read about this second group [Sylviidae] on another page. Gelang et al. (2009) agreed, and proposed the two-family system for babblers. |
Gelang
et al. (2009) also found, based on nuclear and mitochondrial DNA
evidence, that babblers evolved within seven significant clades within
the two major lineages. One big surprise was that one of a half-dozen
genera of wren-babblers — the four species in genus Pnoepygidae
— were not within any babbler lineage. They recommended a new family,
the Pnoepygidae, for those tiny, short-tailed, "wren-babblers." The
remaining genera of birds termed wren-babblers (e.g., Rimator, Ptiolcichla, Kenopia, Napothera, Spelaeornis) are babblers, including the Mountain Wren-Babbler (left in a nice shot by James Eaton), an endemic to the high mountains of northern Borneo. |
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Asia is the center of timalid diversity. Some babblers are tiny and colorful like Chestnut-tailed Minla
(above left), beautifully photographed in n. India by Ron Saldino. They
and other small babblers can be found in mixed species flocks or
isolated pairs inside the forest. Those babblers known as yuhinas (10
species in genus Yuhina) recall North American titmice in their behavior and small crests but many are much more colorful, like Chestnut-crested Yuhina
(above right) in the montane forests of Borneo. But names do not mean
everything. In recent years we have learned that a lot of birds once
considered babblers are not closely related to them at all. Some
examples are:
- As noted above, Sylvia warblers, parrotbills, and Wrentit are babblers, but are now better placed in the "other" babbler family Sylviidae. White-eyes are babblers, and are grouped in this current family. One genera of wren-babbler (Pnoepyga) is not a babbler and deserves it own family.
- White-bellied "Yuhina" is actually a relict vireo living in Asia. It now goes by the name Erpornis Erpornis zantholeuca (Cibois 2003). Shrike-babblers in genus Pteruthius are also relict vireos (Reddy & Cracraft 2007).
- Although
traditionally considered babblers, rockfowl, rockjumpers, and
rockrunners of Africa are not babblers; the first two are now assigned
their own families near the base of the passerine tree, while
rockrunner appears to be in a new "African warbler" lineage that is
also a new family.
- Rail-babbler of tropical
southeast Asia is not a babbler, but is most closely related to
rockjumpers, and is best assigned its own family.
- All the birds named "babblers" in Madagascar, or traditionally listed as babblers in Malagasy field guides, are not babblers.
- Gray-chested "Illadopsis" Kakamega poliothorax, whose generic name commemorates the Kakamega Forest of w. Kenya, is not a babbler but is, along with Spot-throat Modulatrix stictigula and Dapple-throat Arcanator orostruthus, now placed with the sugarbirds in the Promeropidae.
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This
still leaves well over 300 babblers. Asia is the center of babbler
distribution but they also occur in Africa. This family group of Bare-cheeked Babbler
(right) in Namibia were huddled together on one branch, a behavior I've
also seen in other species of babbler in Tanzania and in northern
China. A fair number of species forage together in loose family
groupings.
Before the recent biochemical research,
Fry, Keith & Urban (2000) described the Timaliidae this way: "A
large, heterogeneous, mainly Asiatic family with a complex taxonomic
history and fluctuating composition — many taxa have been moved between
Timaliidae and Sylviidae [Old World warblers] or Turdidae [Thrushes].
Small to quite large passerines, robust, strong-legged, often pale-eyed
... sexes usually alike, rather sombre ... Bills more robust than those
of Sylviidae (probably the closest relatives of babblers)...
Characteristic egg-white protein profile. Syrinx lacks 'turdine thumb'
which is so characteristic of Turdidae and Muscicapidae [Old World
Flycatchers]... Flight weak -- short flights low in dense vegetation,
or of 50m very low over open ground between clumps of shrubs....
Numerous babblers stay in small family parties year-round and are known
or likely co-operative breeders; individuals of many species clump or
huddle tightly together when loafing and roosting...". Some of these
generalities still apply, although genetic evidence now allows us to
clearly separate babblers from thrushes or any of the several families
into which the Old World Warblers have been divided. |
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babblers have a rather subdued plumage pattern although bright rufous
appears in a good many species. Some examples of fairly typical babbler
types scattered among the wide geographic range of babblers include
these: Chestnut-capped Laughingthrush (below left) of montane forests in Borneo, Sumatra & Malaya; Red-billed Leiothrix (below center in a good shot by Ron Saldino) widespread in the Himalayan foothills (and introduced into Hawaii); and Javan Fulvetta (below right) restricted to the mountains of central & western Java. |
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Babblers can be very vocal. Among the Asian babblers are 17 species of fulvettas — another fairly large set of babblers (genus Alcippe)
— all of which have very plain plumage but many of which have lovely
songs given with much verve and energy. Recently I watched a very dull
Brown-cheeked Fulvetta A. poioicephala in Assam, India,
transform itself into a thing of beauty by lustily giving a complex
song from a song perch low in the undergrowth of Panbari Forest. Among
African babblers are the drab but beautifully vocal illadopsis (7
species in the namesake genus) which recall the fulvettas of Asia.
A goodly portion of the babblers are ground-loving species, elusive and
difficult to observe in the forest understory. Some of my favorites are
the wren-babblers — Asia's version of gnateaters. Most of these are
quite small, chubby, long-legged birds with various patterns of spots
or streaks in brown, blackish, or rufescent plumage. There are 20
wren-babblers in 5 genera, almost any one of which is the highlight of
any day's birding. Some are quite specialized (limestone hills, swamps,
bamboo, humid fern forest) and a fair number are rare and local; others
are endemic to Borneo, or the Philippines, or Sumatra. [Note, however,
that one traditional genera of wren-babbler (Pnoepyga) is not within the babbler lineage at all, and is now assigned its own family, Pnoepygidae]. |
Another set of mostly undergrowth species are the 9 species of scimitar-babblers in the genus Pomatorhinus.
These have long decurved bills used to work through the leaf litter —
very like the thrashers of North America which they also resemble in
size and body shape. An example is Rusty-cheeked Scimitar-Babbler
(right) of mountain foothills from India to n. Thailand. Like all birds
of shady habitats, it is difficult to snap a photo without a complex
flash set-up but perhaps you can get an idea of this group from this
white-eyed, pale-billed species. The 7 species of babblers in the genus
Pellorneum are also elusive skulkers. Additional forest
species that can be difficult to locate are mid-canopy species such as
barwings (6 species in Actinodura), sibias (7 species in Heterophasia) and two species of Crocias (Crocias). There are also numerous babblers, some occurring in mixed forest flocks and others as territorial pairs, in the genera Stachyris (28 species), Malacopteron (6 species) and Trichastoma (4 species) to mention just a few others.
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In contrast to the difficult forest babblers, 29 species (10% of all babblers) are open-country birds in the genus Turdoides.
A few are marsh birds but most occur in open savanna or thornscrub
where they travel in small noisy family parties, often leap-frogging
each other as they move quickly from bush to bush so that the whole
party appears to be "rolling" across the plain. The Jungle Babbler
(left) is quite common in the dry forests of the Indian subcontinent;
in Ranthambhor Nat'l Park it has become so tame that it searches the
tourist's jeeps for tidbits. And while the vast majority of babblers
are Asian species, 37 species occur in Africa including 14 of the Turdoides babblers. |
It
is now known that the white-eyes are genetically embedded within the
Babblers; The white-eyes are a mostly homogeneous, Old World tropical
group of small arboreal birds. The name comes not from a white iris,
but from a white-feathered ring that is a prominent feature around a
dark eye of many of these otherwise plain little green birds. Some
standard characters include a brush-tipped tongue for feeding on nectar
(along with insects), semi-pendant nests, and (where known) pale blue
or white eggs (Coates 1990). Some of the white-eyes are montane
specialists, as is the Mountain White-eye (right) on
the Philippines. On some of the larger Philippine islands, three
species of white-eyes divide up elevational zones. Where that occurs,
this species is always at the highest elevation.
Seventy species (74%) of all white-eyes are placed in the genus Zosterops,
and these small birds are often found in chattering flocks moving
through the canopy. Some are common and widespread, but there are a
good many rare or endangered species in the group, including 21 (nearly
a quarter of all species) that were threatened enough to warrant
accounts in Collar et al. (1994). |
White-eyes
are widely distributed in the Old World, with continental
representatives in Africa, Asia, and Australian. However, there are an
inordinate number of island specialties, and no group of birds is such
an "island specialist" (excepting, perhaps, the megapodes). By my
count, 83 white-eyes (88%) are restricted to islands and do not occur
on any major continent, and 70 species (three-quarters of all
white-eyes) occur on only rather small islands (excluding New Guinea,
Borneo, Sumatra, Madagascar, and the larger Philippine islands). There
are two white-eyes endemic to tiny Lifou Island in the Loyalties off
New Caledonia, and many species seem to be described in Clements (1991)
like this for the White-throated White-eye Z. meeki: "Uplands of Tagula
Island (off se New Guinea). Status unknown." No wonder I've seen only a
third of the world's white-eyes!
Many species are isolated on oceanic islands, as is the Caroline Islands White-eye
(left), endemic to Truk (Chuuk), Palau & Pohnpei in Micronesia.
When I first visited Micronesia in 1978, the widespread white-eye over
much of the region was considered a single species, the Bridled
White-eye Z. conspicillatus. Thus, when I photographed this
bird (left) on Truk, it was considered just one of the island races of
Bridled White-eye. Now, however, Bridled White-eye has been split into
at least two species and the bird I photographed is known as the
Caroline Island White-eye (Pratt 1987, Sibley & Monroe 1991).
[There is a special irony associated with that decision and this photo.
Years after I visited Micronesia, I purchased, sight unseen, a booklet
called The Native Forest Birds of Guam (Jenkins 1983). Upon opening it, I was shocked to find that my
photograph of "Bridled White-eye" (the very same bird shown above top
left) was featured in color on the frontispiece of the publication! No
one had asked my permission to do so, but there it was in living color.
At least they spelled my name correctly and did attribute the bird to
Truk.....but how odd that a species that has never occurred on Guam is
the featured photo in the book on Guam.] |
Each
of the islands in the Gulf of Guinea, Africa, has its own endemic, as
do most of the islands in the Indian Ocean. Some of those
representatives, like the Seychelles White-eye Z. modestus
(which I poorly photographed in 1992) have declined during the last
years of the 20th century to endangered status. Others, like the Réunion Olive White-eye (left), are still doing fine.
All
of the white-eyes I've photographed have broad white eye-rings, but a
fair number do not, and sometime go by names such as "Black-eyes."
There is a unique little bird on w. Timor which has been called the
Timor White-eye (Clements 1991), the Spot-breasted White-eye (Sibley
& Monroe 1991), the Spot-breasted Black-eye, and the Timor Ibon.
Barbara Magnuson, who has seen this species with a Ben King tour, tells
me that birds leap-frogged over one another, recalling New World
manakins. It and a related species on Flores are in their own genus (Heleia) and it is possible that these birds are not white-eyes at all. Indeed, Heleia, along with such genera as Rukia in Micronesia, Megazosterops in Palau, Woodfordia in the Solomons, and Hypocryptadius in the Philippines may be another group of birds entirely. Some preliminary DNA work has placed Rukia near Zosterops
(Sibley & Monroe 1991), but most of these taxa have not been
studied. I've seen a couple of them, and could easily believe they were
a family separate and distinct from the otherwise rather homogenous
white-eyes. All lack the broad white eye-rings and none act like
typical white-eyes, and so the word "White-eye" in their name is
misleading. For that reason, Ben King has suggested the Tagalog word
for "small bird" as an appropriate moniker: "Ibon." Thus Hypocryptadius cinnamomeus
becomes the Cinnamon Ibon rather than the misnomer "Cinnamon White-eye"
of Clements (1991). No group deserves a different name more than these
poorly studied, obscure, non-Zosterops "white-eyes." I applaud "Ibon" as a wonderfully coined choice! [Be the first one on your block to use such a cool name....] |
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| Photos: The Streaked Laughingthrush Garrulax lineatus was photographed at Sat Tal in northern India in March 2001. The Silver-eye Zosterops lateralis was eating a crab-apple on Heron I., Great Barrier Reef, Australia, in Sep 1983. James Eaton photographed the Mountain Wren-Babbler Napothera crassa on Mt. Kinabalu, Malaysian Borneo, in June 2002. Ron Saldino photographed the Chestnut-tailed Minla Minla strigula near Lava in w. Bengal, India in late Feb 2001. The Chestnut-crested Yuhina Yuhina everetti was on Mt. Kinabalu, Sabah, Borneo, Malaysia, on 5 Aug 1988, as was the Chestnut-capped Laughingthrush Garrulax mitratus. Ron Saldino photographed the Red-billed Leiothrix Leiothrix lutea near Corbett Nat'l Park, India in mid-March 2001. The Javan Fulvetta Alcippe pyrrhoptera was in Gede Nat'l Park, Java, Indonesia, on 27 Aug 1988. The family of Bare-cheeked Babbler Turdoides gymnogenys was in Erusha Nat'l Park, Namibia, in July 2005. The Rusty-cheeked Scimitar-Babbler Pomatorhinus erythrogenys was at Sat Tal in the Himalayan foothills of Uttar Pradesh, India, in March 2001. The Jungle Babbler Turdoides striatus was in Ranthambhor Nat'l Park, Rajasthan, India, on 23 Mar 2001. The Mountain White-eye Zosterops montanawas on Mt. Kitanglad, Mindanao, Philippines, on 24 Dec 2005. The Caroline Islands White-eye Zosterops semperi was photographed on Moen I., Truk Lagoon, Micronesia, on 31 Aug 1978. The Réunion Olive White-eye Z. olivaceus was in the forest above Le Brulé, Réunion I., Indian Ocean, on 4 Dec 1992. All
photos © Don Roberson except those attributed to Ron Saldino or
James Eaton, respectively, who hold those copyrights (used with
permission here); all rights reserved. |
Bibliographic notes
There
is no recent "family book" covering the Babblers but good recent
coverage of African species is in Fry et al. (2000), and the Handbook of the Birds of the World
account (Collar & Robson 2007) is wonderful, with exceptional
photos, although it could not have been quite up with the most current
taxonomic findings.
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.
Birdlife International. 2000. Threatened Birds of the World. Lynx Edicions, Barcelona, Spain.
Cibois, A. 2003. Mitochondrial DNA phylogeny
of babblers (Timaliidae). Auk 120: 35-54.
Cibois,
A., E. Pasquet, and T.S. Schulenberg. 1999. Molecular systematics of
the Malagasy babblers (Timaliidae) and Warblers (Sylviidae), based on
cytochrome b and 16S rRNA sequences. Molec. Phylog. Evol. 3: 581-595.
Cibois,
A., B. Slikas, T.S. Schulenberg, and E. Pasquet. 2001. An endemic
radiation of Malagasy songbirds is revealed by mitochondrial DNA
sequence data. Evolution 55: 1198-1206.
Cibois, A. 2003. Mitochondrial DNA phylogeny of babblers (Timaliidae) Auk 120: 35-54.
Clements, J.F. 1991. Birds of the World: A Check-List. 4th ed. Ibis Publishing, Vista, CA.
Coates, B. J. 1990. The Birds of Papua New Guinea. Part II. Dove Publ., Ltd., Alderley, Australia.
Collar,
N.J., M.J. Crosby, and A. J. Stattersfield. 1994. Birds to Watch 2: The
World List of Threatened Birds. Birdlife Conserv. Ser. 4. Birdlife
Intern'l, Smithsonian Inst., Washington, D.C.
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.
Dickinson,
E.C., ed. 2003. The Howard & Moore Complete Checklist of the Birds
of the World. 3d ed. Princeton Univ. Press, Princeton, N.J.
Fry, C. H., and Keith, S., and E. Urban., eds. 2000. The Birds of Africa. Vol. VI. Academic Press, London.
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.
Grimmett, R., C. Inskipp, and T. Inskipp. 1998. Birds of the Indian Subcontinent. Christopher Helm, London.
Jenkins, J. M. 1983. The Native Forest Birds of Guam. Ornithol. Monogr. 31. A.O.U., Washington, D.C.
Jønsson,
K.A., and J. Fjeldså. 2006. A phylogenetic supertree of oscine
passerine birds. Zoologica Scripta 35: 149-186.
Keith, S., E. Urban, and C. H. Fry, eds. 1992. The Birds of Africa. Vol. IV. Academic Press, London.
MacKinnon, J., and K. Phillipps. 2000. A Field Guide to the Birds of China. Oxford Univ. Press, Oxford.
Pratt,
H. D., P. L. Bruner, and D. G. Berrett. A Field Guide to the Birds of
Hawaii and the Tropical Pacific. 1987. Princeton Univ. Press,
Princeton, N. J.
Reddy, S., and J. Cracraft. 2007. Old World Shrike-babblers (Pteruthius) belong with New World Vireos (Vireonidae). Molec. Phylog. Evol. 44: 1352-1357.
Sibley,
C. G., and J. E. Ahlquist. 1990. Phylogeny and Classification of Birds:
A Study in Molecular Evolution. Yale Univ. Press, New Haven, CT.
Sibley, C. G., and B. L. Monroe. 1990. Distribution and Taxonomy of Birds of the World. Yale Univ. Press, New Haven, CT.
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