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MONARCHS Monarchidae
Monarch flycatchers, Paradise-Flycatchers, and Magpie-Larks |
- 89 species in the Old World tropics and on Pacific & Indian Ocean islands
- DR personal total: 33 species (35%), 10 photo'd
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The
Monarchs are a rather large Old World family of (mostly) arboreal
flycatchers. Like the New World flycatchers (Tyrannidae) and the Old
World flycatchers (Muscicapidae), many of them forage for flying
insects by sallying out from perches, and many build cup-like nests.
Many have previously been called "flycatchers," but the modern trend is
to call them "monarchs," as they are not related to the other
"flycatcher" families.
The Monarchs reach their
greatest diversity in the Old World tropics on islands between Asia and
Australia. Some very dramatic birds occur in the Philippines,
Indonesia, and New Guinea. One is Rufous-collared Monarch (left, on a nest in a fine photo by Steve Wilson). It is one of the monarchs to show colorful bare wattles around the eye.
Although monarchs are not large in size, some of their plumage features are very impressive. Celestial Monarch
(below left, in an amazing shot by Mark Sutton), in the lowlands of a
couple Philippine islands, has a electric sky-blue crest,. White-naped Monarch (below right), endemic to the northern Moluccas, is black-and-white with amazing facial tufts. |
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Many
of these wonderful birds are in decline; some are endangered. Celestial
Monarch has been recently found at only 10 sites in its severely
fragmented lowland habitat (Birdlife International 2000). During my
first visit to the Philippines in 1990, no one knew where or how to
find Celestial Monarch. It was a mythical bird. By 2005, though, the
late Tim Fisher and his cadre of local guides knew how to find it: one
must know and listen for its call and then track it down in the canopy.
This can be exceedingly difficult to do. We spent many hours in the
muggy lowland forest searching for this bird. When it was found, not
all of our group got on it. It just disappears high in the canopy. But
those of us who did see it learned, as shown in Sutton's photo, that
the crest lies flat atop the head. It is listed as Endangered. |
Among endangered monarchs, Cerulean Paradise-Flycatcher Eutrichomyias rowleyi,
restricted to the island of Sangihe, Indonesia, was considered extinct
by Clements (1991), but was rediscovered during surveys in 1998 (but
just 21 birds at six sites; Birdlife International 2000). Many endemic
monarchs to small Wallacean islands have also been rediscovered this
past decade, and for some populations are better than had been feared.
Others remain problematic. Some 15 species of monarchs from Pacific
Ocean islands are listed as seriously threatened with extinction
(Birdlife International 2000). Those in the "critical" category include
Tahiti Monarch Pomarea nigra (just 9 pairs and 24 birds remaining as of Jan 2000), and Fatuhiva Monarch P. whitneyi (endemic to Fatu Hiva in the Marquesas in density of only 1 pair/10 ha).
Taxonomy is also a problem and has been in flux for some time. Take Spectacled Monarch
for example. There is a complex of populations from Halmahera and the
Lesser Sundas to eastern Australia. As explained by Rheindt &
Hutchinson (2007), those population that have lost rufous coloration to
the breast and neck have been elevated to species status — such as
Black-chinned Monarch S. boanensis and Black-tipped Monarch S. loricatus —
leaving between "a swiss cheese" complex of rufous-breasted monarchs
from Indonesia to Australia, all called Spectacled Monarch S.
trivirgatus, even though Black-chinned Monarch is considered a "good"
species that exists on a tiny island just a few miles of Seram, which
has a rufous-breasted bird. Clearly a thorough review and revision is
necessary. As it currently stands, some world checklists (such as IOC)
split the Halmahera bird (shown above) as "Moluccan Monarch S. bimaculatus"
This may be proper but it is very piecemeal, since the entire complex
needs revision. ["Moluccan Monarch" is also a poor English name, since
other Moluccan populations likely are of species-level status; see
Rheindt & Hutchinson (2007) on the Seram population.] Filardi &
Moyle (2005) showed that the genus Monarcha was paraphyletic and had to be split into at least 3 genera; Symposiachrus was resurrected for Spectacled Monarch and close relatives. Then Filardi & Smith (2005) focused in on the Symposiachrus
monarchs, and found that Spectacled Monarch was paraphyletic as a
species, since the resident populations on Cape York, Australia, were
more closely related to Spot-winged Monarch S. guttula of New Guinea
than they were to migrant population of Spectacled Monarch elsewhere in
eastern Australia (S. t. gouldi). A lot more work is needed. |
Beyond
the intriguing monarchs of the Pacific tropics, though, the family
Monarchidae has other fabulous birds. Among the loveliest are 15
paradise-flycatchers in two genera. Most are in the genus Terpsiphone, like this Red-bellied Paradise-Flycatcher
(left). The species frequents the dense forest of the Congo Basin and
west Africa; here it is on a nest then under construction in Uganda.
Other species are migratory, such as the Asian Paradise-Flycatcher T. paradisi.
Its annual arrival at Bharatpur in north-central India heralds the
beginning of spring there, just as American Robins do in the eastern
United States.
There are some species of paradise-flycatcher that have dramatic sexual dimorphism. A good example is the Seychelles Paradise-Flycatcher
(below), a rare and threatened species currently restricted to a single
island (La Digue) in the Seychelles. The female (shown here on a nest)
is tricolored in black, rust, and white; she has only a medium length
tail for a paradise-flycatcher. The all-black male, however, has an
exceptionally long tail that drags behind, undulating in flight. |
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Paradise-Flycatchers are known for their beautiful long tails in adult males. Madagascar Paradise-Flycatcher (right, in a nice shot by Murray Lord). Not all species of paradise-flycatchers have that trait. The Mascarene Paradise-Flycatcher
(below), for example, has rather medium length tails in both sexes.
This photo is of a female; she, like almost all members of the
Monarchidae, has a rather shallow, broad bill. Monarchs are often
characterized by broad, flattened bills with a slightly hooked tip, a
subterminal maxillary notch, rictal bristles and rounded nostrils
(Sibley 1996).
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Africa
may the center of the paradise-flycatcher world, with 7 of 15 species,
but I find it particularly interesting that ancestral flycatchers made
it to offshore islands in the Indian Ocean and evolved into separate
species on Madagascar [Terpsiphone mutata], on Mauritius and Réunion [T. bourbonnensis], on the Seychelles [T. corvina], and on Annobón Island [T. smithii]. There is also a species on Sao Tomé Island in the Gulf of Guinea off west Africa [T. atrochalybeia].
There are also paradise-flycatchers in the western Pacific. A species in Japan migrates to the Asian mainland in winter [T. atrocaudata], and there are two species on the Philippines. Rufous Paradise-Flycatcher
(right, in a lovely photo by Blake Matheson) lives in humid lowland
forests. These habitats are rapidly being lost on Mindanao, where this
picture was taken. Despite its brilliant color, one primarily finds
Rufous Paradise-Flycatcher by voice. The jungles are just that thick
and difficult. |
Sibley
& Ahlquist (1990) showed that the monarchs were among those evolved
from the great corvid assemblage that arose in Australasia, and
certainly the family's center of abundance is there and stretching
westward to Wallacea and the Philippines. Filardi & Moyle (2005)
showed, though, that the spread of monarchs was not simply
"downstream," i.e., from continents to islands. Through molecular
evidence they found that some "upstream" colonization occurred in
monarchs — members of island populations later colonized continental
Australia and diversified.
Outside of the
'mainstream' Monarchidae, much of it on islands, and aside from the
radiation of paradise-flycatchers in Africa and Asia, rather few
monarchs have colonized and survived in mainland Asia. One is the
widespread Black-naped Monarch Hypothymis azurea which ranges from India to Borneo and western Indonesia. Recent split from that species is Pale-blue Monarch (left), an endemic in the Sulawesi lowlands.
There
are also 7 or so monarchs, plus Magpie-lark (see below), that breed in
continental Australia. Like many in the family, they build
tightly-woven open-cup nests. This Restless Flycatcher (below) is sitting tight on such a nest. |
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A
most impressive example of distributional radiation and evolution are
the many species of monarchs far-flung across the Pacific. There are
shrikebills (Clytorhynchus) on Fiji, New Caledonia, and the Solomons. There are monarchs on many islands in Melanesia and Micronesia.
One monarch even reached Hawaii: the 'Elepaio
(left), which then evolved subspecific differences on three islands
(Kauai, Oahu, Hawaii), including three subspecies on the big island of
Hawaii itself (Pratt et al. 1987). The 'Elepaio shown is the race on the Hilo side of Hawaii C. s. ridgwayi, called the "Volcano 'Elepaio".
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The saddest part of the Pacific island story is that quite a number of species are now threatened with extinction. Although the Myiagra
monarchs on some islands are still common, others are losing ground
rapidly and at least one is already lost. In 1979 a trip around the
world with my father and sister took me to Guam. With just one day to
go birding, that I saw the Guam Flycatcher Myiagra freycineti
(also known as Guam Myiagra or, back then, Guam Boatbill. It was in the
remnant forest at the north tip of Guam. That was 29 Aug 1979, and I
wrote a brief field description of the bird. It was then thought to be
fairly common. A horrendous population crash of all Guam birds,
including the monarch, took place in the 1980s, primarily due to the
accidental introduction of the Brown Tree Snake Boiga irregularis
from New Guinea. There were no native snakes on Guam. This non-native
predator, against which birds had evolved no defensive strategies,
essentially wiped out the native landbirds of Guam . Some, like the
Guam Rail Galliralus owstoni, were saved by capturing
remaining rails for captive breeding (I saw a wild Guam Rail the same
day as the Guam Monarch, before the population collapses). But the
flycatcher was decimated. It was extinct within the decade. Just 30
years ago I saw that lovely little monarch as a wild bird. Today it is
extinct, never to be seen again. It is hard to accept things like that.
There are still an assortment of Myiagra flycatchers in Australasia and the south Pacific. One is Moluccan Flycatcher
(or Slaty Myiagra; below), endemic to the Moluccas, and it is dimorphic
in plumage. The female (below mddile) has a pattern rather similar to
the now-extinct Guam Flycatcher — grayish above with a buffy wash to
the breast. The male Guam Flycatcher was glossy blue-black and also had
a buffy wash to breast; male Moluccan (below left) is similar but is
white-breasted. Many of the Myiagra flycatchers have this
generalized pattern. On the final photo (below right) we can see the
bill shape of the male Moluccan –— and similar to other Myiagra flycatchers — that gave rise to the old name "boatbill" for these little birds. The bill is very wide. |
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On a brighter side, one final set of monarchs appear to be doing fine. Indeed, Magpie-Lark (left) is common in Australia and southern New Guinea. It, and the Torrent-lark Grallina bruijni
in montane New Guinea, have traditionally been considered to be the two
members of the family Grallinidae. However, molecular evidence
indicates that the two members of the genus Grallina are
deeply embedded within the monarch assemblage (Baverstock et al. 1992,
Sibley 1996, Schodde & Mason 1999). Unfortunately, the assignments
of text and species in the Handbook of Birds of the World
series were, of necessity, decided years prior to publication. The
initial decision was to treat the Grallinidae as a family. By the time
of the publication of the Monarchidae (Coates et al. 2006) it was
acknowledged that the "most recent analysis strongly indicates that the
magpie-larks are, indeed, monarchs" (p. 247). Yet it was too late to
change, and "Grallinidae" has a family chapter in Vol. 14. |
Schodde
& Mason (1999) wrote, as to revising the placement of magpie-larks
into the clade with drongos, fantails and monarchs: "Molecular and
cranial studies have since shown that magpie-larks are unquestionably
dicrurids. This makes sense of their slow buoyant flight, colour
pattern and sexual dimorphism which is Monarcha-like; even pallid bill, inner nest structure, egg markings and the Myiagra-like
begging calls of its fledglings fit." These words make even better
sense when placing them directly in the Monarchidae itself. Indeed,
they are so embedded that most recent authors do not even give them
subfamily status (e.g., Dickinson 2003).
The distinctively patterned Magpie-lark
(right) is widespread in Australia (Blakers et al. 1984). It is often
one of the first new birds seen on a visit "down under." They walk in
pastures and cleared areas wherever water is nearby, including the
lawns on the Cairns' esplanade (above left). They also reside in the
drier plains of the interior, where this adult was carrying food for
nestlings (right).
The other species in the genus,
Torrent-lark, occurs only in montane New Guinea where it haunts rocky,
swift-flowing streams. It forages on exposed rocks in streams, or
sometime wet gravel nearby (even on roads; Coates 1990). This sounds
like the niche filled by the forktails in southeast Asia. |
The
Magpie-lark builds thick cup-like mud nests; fledged young birds (left)
are fed by the parents for a time, and then form erratic flocks. This
is the closest to any wandering made by most of these birds, although
some seasonal movements have been noted in the Torres Strait region and
vagrants do occasionally occur to Tasmania (Blakers et al. 1984). Birds
mostly occur in pairs or small family parties.
Clements
(1991) uses the term "Mudnest Builders" for the Grallinidae but some
Australian texts (e.g., Simpson & Day 1996) calls the Corcoracidae
(Apostlebirds) the "Australian Mud-Nesters" [species of both groups
build mud nests]. They were once considered part of the same family but
they are unrelated. The building of mud nests was just a common
adaptation to living in open interior country — the same adaptation is
made by the horneros in South America. |
Finally,
there are some birds of ancient lineage in the corvoid assemblage that
may be basal elements of the Monarch evolutionary tree. The Blue-capped Ifrita Ifrita kowaldi
may be an early branch of monarchs, and best assigned to this family.
This tentative hypothesis is suggested by molecular studies (e.g.,
Barker et al. 2004, Norman et al. 2009, Jønsson et al. 2011) but
the Ifrita remains an enigma of uncertain relationships. I have
separate pages on this mystery bird at the link at its name. |
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Photos: Steve Wilson photographed the Rufous-collared Monarch Arses insularis
at a nest behind the research center at Baiyer River reserve, Papua New
Guinea, on 27 Oct 1983 (my photo attempt didn't work on this bird).
Mark Sutton photographed the Celestial Monarch Hypothymis coelestis at Apaya, Luzon, Philippines, on 6 Apr 2004. The White-naped Monarch Carterornis pileatus was at Foli, Halmahera, Indonesia, on 7 Oct 2011. The Halmahera race (species?) of the Spectacled Monarch Symposiachrus trivirgatus bimaculatus was at the mountain pass on the Buli Road in Halmahera, Indonesia, in Oct 2011. The Red-bellied (sometimes called Black-headed) Paradise-Flycatcher Terpsiphone rufiventer was photographed at Kibale Forest, Uganda, 26 July 2002. Murray Lord photographed the Madagascar Paradise-Flycatcher Terpsiphone mutata near Ifaty, Madagascar, on 8 October 2005. The Mascarene Paradise-Flycatcher Terpsiphone bourbonnensis was above Le Brulé, Réunion Island, on 4 Dec 1992. The Seychelles Paradise-Flycatcher Terpsiphone corvina was on La Digue I., Seychelles, on 13 Nov 1992. Blake Matheson photographed the Rufous Paradise-Flycatcher Terpsiphone cinnamomea in the PICOP forest, Mindanao, the Philippines, on 28 Dec 2005. The Pale-blue Monarch Hypothymis puella was at Tangkoko NP, Sulawesi, Indonesia, in Oct 2011. The nesting Restless Flycatcher Myiagra inquieta was in the Hawkesbury Valley, N.S.W., Australia, on 18 Sep 1983. The 'Elepaio Chasiempis sandwichensis was in the Hakalau Forest NWR, Hawaii, Hawaii, on 2 Jan 2012. The Moluccan Flycatcher Myiagra galeata was at Foli, Halmahera, Indonesia, on 7 Oct 2011. The Magpie-lark Grallina cyanoleuca
photos were taken, respectively: at Cairns, Queensland, in Nov 1983;
carrying food at Deniliquin, New South Wales, on 1 Jan 1998; and the
still dependent fledgling near Mareeba, Queensland, in Jan 1998. Photos
© Don Roberson, except Rufous-collared Monarch © Steve Wilson, Celestial Monarch © Mark Sutton, and Rufous Paradise-Flycatcher © Blake Matheson, all used with permission; all rights reserved.
Bibliographic note: There is no family book but the Handbook of the Birds of the World
chapter (Coates et al. 2006) is a good introduction and has many superb
photographs, although it does include various birds that are now known
not to be monarchs, and fails to include Magpie-lark and Torrent-lark,
which are. Additional great photographs of many New Guinea monarchs are
in Coates (1990).
Literature cited:
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.
Baverstock,
P.R., R. Schodde, L. Christidis, M. Krieg, and J. Birrell. 1992.
Evolutionary relationships of the Australasian mud-nesters
(Grallinidae, Corcoracidae): immunological evidence. Australian J.
Zool. 40: 173–179.
Beresford, P., F.K. Barker, P.G. Ryan, and
T.M. Crowe. 2005. African endemics span the tree of songbirds
(Passeri): molecular systematics of several evolutionary 'enigmas'.
Proc. R. Soc. B 272: 849–858.
Birdlife
International. 2000. Threatened Birds of the World. Barcelona &
Cambridge, U.K., Lynx Edicions & Birdlife International.
Blakers,
M., S.J.J.F. Davies, and P.N. Reilly. 1984. The Atlas of Australian
Birds. Royal Australian Ornith. Union, Melbourne Univ. Press, Carlton,
Victoria.
Clements, J.F. 1991. Birds of the World: A Checklist. 4th ed. Ibis Publ., Vista, CA.
Coates, B.J. 1990. The Birds of Papua New Guinea. Part II. Dove Publ., Ltd., Alderley, Australia.
Coates, B.J., G.C.L. Dutson, and C.E. Filardi. 2006. Family Monarchidae (Monarchs), pp. 244–329 in Handbook of the Birds of the World (J. del Hoyo, A. Elliott, & D.A. Christie, eds.). Vol. 11. Lynx Edicions, Barcelona.
Dickinson, E., ed. 2003. The Howard & Moore Complete Checklist of
the Birds of the World. 3d ed. Princeton Univ. Press, Princeton, N.J.
Filardi,
C.E., and R.G. Moyle. 2005. Single origin of a pan-Pacific bird group
and upstream colonization of Australasia. Nature 438: 216-219.
Filardi, C. E., and C. E. Smith. 2005. Molecular phylogenetics of monarch flycatchers (genus Monarcha) with emphasis on Solomon Island endemics. Molec. Phylog. Evol. 37: 776–778.
Jønsson,
K.A., F. Pierre-Herni, R.E. Ricklefs, and J. Fjeldså. 2011. Major
global radiation of corvoid birds originated in the proto-Papuan
archipelago. Proc. Nat. Acad. Sci. 108: 2328-2333.
Norman,
J.A., P.G.P. Ericson, K.A. Jønsson, J. Fjeldså, and L.
Christidis. 2009. A multi-gene phylogeny reveals novel relationships
for aberrant genera of Australo-Papuan core Corvoidea and polyphyly of
the Pachycephalidae and Psophodidae (Aves: Passeriformes). Molec.
Phylog. Evol. 52: 488-497.
Pratt, H. D., P. L.
Bruner, and D. G. Berrett. 1987. A Field Guide to the Birds of Hawaii
and the Tropical Pacific. Princeton Univ. Press, Princeton, N.J.
Rheindt,
F.E. and R. Hutchinson. 2007. A photoshot odyssey through the confused
avian taxonomy of Seram and Buru (southern Moluccas). Birding Asia 7:
18–38.
Schodde, R., and I.J. Mason. 1999. The Directory of Australian Birds, Passerines. CSIRO Publishing.
Sibley, C. G. 1996. Birds of the World, on diskette, Windows version 2.0. Charles G. Sibley, Santa Rosa, CA.
Sibley, C. G., and J. E. Ahlquist. 1990. Phylogeny and Classification
of Birds: A Study in Molecular Evolution. Yale Univ. Press, New Haven,
CT.
Simpson, K, and N. Day. 1996. A Field Guide to
the Birds of Australia, rev. 5th ed. Penguin Books Australia Ltd.,
Ringwood, Victoria, Australia.
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