Upload
others
View
3
Download
0
Embed Size (px)
Citation preview
See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/273450387
Return to the Malay Archipelago: the biogeography of Sundaic rainforest birds
Article · March 2015
DOI: 10.1007/s10336-015-1188-3
CITATIONS
26READS
523
3 authors:
Some of the authors of this publication are also working on these related projects:
Birds Fiji View project
oVert Thematic Collections Network View project
Frederick H Sheldon
Louisiana State University
154 PUBLICATIONS 6,005 CITATIONS
SEE PROFILE
Haw Chuan Lim
George Mason University
40 PUBLICATIONS 899 CITATIONS
SEE PROFILE
Robert Glen Moyle
University of Kansas
208 PUBLICATIONS 3,122 CITATIONS
SEE PROFILE
All content following this page was uploaded by Frederick H Sheldon on 18 December 2018.
The user has requested enhancement of the downloaded file.
REVIEW
Return to the Malay Archipelago: the biogeography of Sundaicrainforest birds
Frederick H. Sheldon • Haw Chuan Lim •
Robert G. Moyle
Received: 12 November 2014 / Revised: 11 February 2015 / Accepted: 23 February 2015
� Dt. Ornithologen-Gesellschaft e.V. 2015
Abstract During the last 15–20 years, phylogenetic,
phylogeographic, paleontological, geological, and habitat
modeling studies have improved our knowledge of Sundaic
biogeography dramatically. In light of these advances, we
review (or postulate) where Sundaic rainforest birds came
from, the causes of their endemism, and the influence of
Pleistocene climatic perturbations on their diversification.
We suggest that four scenarios make up a coherent, plau-
sible explanation of patterns of extant diversity. First, re-
lictual lineages, which represent hangovers from the warm,
wet Eocene, survived the hard climatic times of the colder,
drier Oligocene and Pliocene in the mountains and adjacent
lowlands of eastern Borneo, where rainforest has existed
continuously for the last 20–30 million years. Second, most
modern SE Asian genera developed during the Miocene.
Third, the rainforest of Sundaland and its avifauna were
largely isolated from the rest of SE Asia during the late
Miocene and Pliocene by seasonal habitats in southern
Indochina and ocean boundaries elsewhere, increasing re-
gional endemism. Finally, the advent of global glaciation in
the Pleistocene introduced a different diversification dy-
namic to Sundaland. Early glacial events caused sufficient
drying in central Sundaland to fragment rainforest and its
avifauna into refugia in eastern and western Sundaland and
to allow dry-habitat taxa to reach Java from Indochina.
More recent glacial events resulted in sufficient perhumid
habitat in central Sundaland to reconnect previously vi-
cariated rainforest populations, creating the lowland and
elevational parapatry we see today. This Pleistocene dy-
namic was probably not simply one period of separation
and one period of connection, but rather a complex inter-
play of isolation and colonization, influenced by highly
variable population sizes, changing levels of gene flow, and
behavioral idiosyncrasies of the species involved.
Throughout all of these events, Borneo played a seminal
role in rainforest bird evolution by providing the habitat
necessary for diversification and the long-term survival of
taxa.
Keywords Avifauna � Borneo � Pleistocene glaciation �Java � Savanna � Sundaland
Introduction
The diversity and distribution of Indo-Malayan birds in-
spired Alfred Russel Wallace as he labored in the forests
and bungalows of Sarawak 160 years ago (Wallace 1876,
1883), and they still intrigue us today. Where did the Rail-
babbler (Eupetes macrocerus), Bornean Bristlehead (Pi-
tyriasis gymnocephala), and other unusual species of the
Greater Sundas come from? Why are so many Sundaic
endemics montane? Why are almost all SE Asian trogons
Communicated by E. Matthysen.
Electronic supplementary material The online version of thisarticle (doi:10.1007/s10336-015-1188-3) contains supplementarymaterial, which is available to authorized users.
F. H. Sheldon (&)
Museum of Natural Science and Department of Biological
Sciences, Louisiana State University, Baton Rouge, LA, USA
e-mail: [email protected]
H. C. Lim
Smithsonian Institution, National Museum of Natural History,
Washington, DC, USA
R. G. Moyle
Biodiversity Institute and Department of Ecology
and Evolutionary Biology, University of Kansas,
Lawrence, KS, USA
123
J Ornithol
DOI 10.1007/s10336-015-1188-3
uniform in morphology and voice (Harpactes), except for
one taxon in the mountains of Java and Sumatra (Apal-
harpactes)? Why do barbets exhibit much the same pattern:
30 species of Megalaima (including Psilopogon)—almost
all with uniformly green bodies and colorful heads,
monotonous ‘‘tooking’’ or ‘‘pooping’’ songs, and solitary
lifestyles—versus one odd brown, wheezing, highly social
taxon (Caloramphus)? Why does Java share some taxa
with Indochina but not with Sumatra and Borneo? Why are
Bornean populations subdivided taxonomically despite
continuous rainforest across the island, such that species
like the Black-thighed Falconet (Microhierax fringillarius)
and Garnet Pitta (Pitta granatina), which inhabit most of
Borneo, are suddenly replaced in the northeast by the
White-fronted Falconet (M. latifrons) and Black-crowned
Pitta (P. ussheri)?
In this review, we try to come to grips with these and
other enigmatic patterns evident in the rainforest avifauna
of the Greater Sundas. The tremendous amount of phylo-
genetic, geological, geographic, and paleontological in-
formation generated in the last 15–20 years makes it
possible to propose with some degree of confidence: (1)
where major components of the Sundaic avifauna
originated on a global scale, (2) roughly (very roughly)
when they arrived in SE Asia, (3) what caused major
groups to radiate or decline in the Sunda Islands, and (4)
how diversification occurred in more recent geological
times. Our examples come mainly from Borneo because we
are most familiar with Bornean birds, and Borneo is by far
the best-studied Sundaic landmass in terms of phyloge-
netics and population genetics. We also discuss bird evo-
lution on the other Sunda islands, particularly Java,
because the unique histories of these islands help illumi-
nate biogeographic patterns. What is evident in this telling
is that we have a reasonable idea where some groups of
Indo-Malayan birds ultimately came from and what forces
likely caused Pleistocene diversification—but much of our
understanding of Sundaic bird evolution is shaky at best.
Dates attributed to events, for example, may be off by tens
of millions of years in some instances. Regardless of this
substantial uncertainty, we present a fairly simple, in some
cases provocative, outline of biogeographic history in the
hope of stimulating more rigorous investigations of plau-
sible scenarios in the future.
Major patterns in the Sundaic avifauna may be attributed
to just a few interrelated climatic and geographic forces,
the most important of which were: (1) changes in tem-
perature and precipitation between Cenozoic epochs, (2)
climatic changes associated with global glacial events of
the Pleistocene, (3) persistence of Sundaic rainforest
refuges through hard climatic times, and (4) the topography
and position of Borneo. Climatic shifts between Cenozoic
epochs (Berggren and Prothero 1992; Zachos et al. 2001)
almost certainly fostered cycles of taxic radiation and ex-
tinction in SE Asia (Morley 2000; Mittelbach et al. 2007),
sort of ‘‘taxon cycles’’ written on a large scale (Ricklefs
and Cox 1978). The warm wet Eocene (a hot house epoch)
would have led to a bloom of rainforest species, the cold
dry Oligocene (an ice house epoch) to a reduction in
rainforest species, the warm wet Miocene to a bloom, and
the cold dry Pliocene to a reduction. The Eocene bloom
and Oligocene reduction are suggested by the age and
characteristics of a few rainforest relicts that have survived
to modern times, and the Miocene bloom is evident from
phylogenetic studies of modern taxa. The persistence of
rainforest refuges in and around mountains would have
allowed rainforest species to survive through the cooler,
drier Oligocene and Pliocene. The Pleistocene Ice Age
began at about 2.6 Ma (Mega-annum or million years ago),
and subsequent global glacial and interglacial events in-
troduced a new dimension to the process of diversification
in Sundaland. Habitat alterations wrought by sea level and
climate changes during the glacial events explain much of
the parapatry we see today between populations both
across islands in lowland rainforest and elevationally on
mountains. They also explain the disjunction between Ja-
van and Indochinese taxa. Throughout much of the Ceno-
zoic, Borneo must have played a special role in rainforest
bird diversification and preservation because of its age,
size, extensive mountains, and position on the eastern edge
of Sundaland (de Bruyn et al. 2014). In contrast, the other
Greater Sunda Islands are relatively recent derivatives. As
such, these islands would have inherited much of their
rainforest avifauna from Borneo and to a lesser extent the
Malay Peninsula, which did not enjoy the rainfall or well-
positioned mountains necessary to preserve extensive
rainforest through colder, drier times.
Our review begins with traditional biogeographic de-
scriptions of Sundaland and its avifauna. We then scan the
geological and geographic history of the region from
60–5 Ma, highlighting types of land masses, habitats, and
bird groups likely to have existed in SE Asia at various
points in time. Starting at 5 Ma, in the Pliocene, the pace of
the review slows, and discussion focuses on geographic
changes, especially those driven by global glacial events of
the Pleistocene Ice Age, and how they influenced species
distributions, population structure, and diversification in
general.
The geography and avifauna of Sundaland
The Malay Peninsula, Borneo, Sumatra, Java, Palawan, and
many smaller islands lie on the Sunda Continental Shelf
(Fig. 1), and together they constitute Sundaland (‘‘Sunda
Land’’; Molengraaff 1921). This is an important
J Ornithol
123
biogeographic subregion of SE Asia because of its
relatively non-seasonal (perhumid) rainforest and long-
term existence. To its north, Sundaland is separated from
Indochina by the transition from rainforest to more sea-
sonal forest. The floral demarcation of this shift is the
Kangar–Pattani Line (van Steenis 1950), but most verte-
brate biologists associate the Sundaic border with the
Isthmus of Kra further to the north, at approximately 10�latitude (Woodruff and Turner 2009). Biogeographic issues
related to this transition were reviewed by Hughes et al.
(2003) and Round et al. (2003). To the east, Sundaland is
separated from Wallacea by the deep oceanic trench of the
Makassar Strait, along which runs Wallace’s Line (Whit-
more 1981; Lohman et al. 2011). To the northeast, Sun-
daland is separated from the oceanic Philippine islands by
Huxley’s Line (Esselstyn et al. 2010).
Sundaland shares the largest proportion of its resident
forest birds with Indochina, and fewer with the Philippines
and Wallacea (Mayr 1944; Darlington 1957). Despite this
commonality, much of Sundaland’s avifauna is distinct
because of the habitat and physical barriers that have iso-
lated it for millions of years. About 691 resident land bird
species inhabit Sundaland, of which 264 (38 %) are en-
demic to the region [Table 1; Electronic Supplementary
Material (ESM) 1]. Among the Sundaic landmasses, the
Malay Peninsula has the most species (420) and the fewest
endemics (4). This odd numerical combination stems from
the Peninsula’s position, which results in it sharing large
numbers of species with Indochina and the Sunda Islands
(Wells 1999). Sumatra has the most species of any of the
Sunda islands (414), but not the most endemics (33), even
though it includes the endemic-rich Mentawai and
Enggano islands to its southwest. Borneo ranks third in
terms of number of species (373), but it has the most en-
demics (52), most of which are montane (Smythies 1999;
Phillipps and Phillipps 2014). Borneo also has the most
frogmouths, trogons, hornbills, barbets, pittas, broadbills,
flowerpeckers, and spiderhunters of any place in the world
(Phillipps and Phillipps 2014). Palawan possesses the
fewest species of any major Sunda island, presumably
because of its small size and relative isolation (Esselstyn
et al. 2010; Lim et al. 2014). Java has the most distinct
avifauna of any Sunda island by virtue of its position next
to the Lesser Sundas and its drier and more seasonal
Fig. 1 Map of Sundaland, with
the Greater Sunda Islands and
outlying biogeographic regions
(Indochina, Philippines, and
Wallacea) indicated. Black lines
Borders of Sundaland
Table 1 Total number of resident and endemic land bird species in
Sundaland
Area Total species (n) Endemics (n) % Endemism
Malay Peninsula 420 4 1
Borneo 373 52 14
Sumatra 414 33 8
Java 313 41 13
Palawan 153 18 12
Sundaland 691 264 38
Based on the list of Sundaic species in the ESM 1 and the classifi-
cation of Gill and Donsker (2014)
J Ornithol
123
lowlands (Whitten et al. 1996). Java’s avifauna also com-
prises about 30 species with disjunct relationships to taxa
in Indochina (MacKinnon and Phillipps 1999).
The Malay Peninsula and Sumatra share the most resi-
dent land bird species (342) of any of the Sundaic land-
masses (Table 2), as would be expected by their proximity
and parallel geographic positions. These two areas are most
similar to Borneo in habitat and size and, thus, share the
most species with that island, about 280 and 289, respec-
tively. Java shares about 200 species each with the Malay
Peninsula, Borneo, and Sumatra, and Palawan shares about
85 species with each of the other areas.
From the Eocene to the Pliocene
Biogeography of the Indian Ocean
The birds of Sundaland come not only from adjacent Asia
but also from India, Africa, and Australia (Fig. 2). Re-
constructing the evolutionary history of Sundaland thus
requires an understanding of the Cenozoic geography of
the entire Indian Ocean region.
At the beginning of the Cenozoic, at around 66 Ma,
Sundaland was a peninsula jutting south from SE Asia
(Fig. 3), and its flora and fauna were presumably Laur-
asian. India was the first landmass with the potential to
bring Gondwanan land birds to Laurasia, at between 50 and
35 Ma (Morley 1998b; Aitchison et al. 2007; Kumar et al.
2007; Morley 2012; Hall 2013; Li et al. 2013). The timing
of India’s collision with Asia is not known precisely.
However, any exchange of organisms between India and
SE Asia would have been complicated by the development
of land bridges before complete continental unification and
the habitat barriers that followed unification (Hall 2013),
not to mention overwater dispersal. Indian flora arrived in
Sundaland in the mid-Eocene, suggesting a moist corridor
between the two regions (Morley 1998b, 2012). Whether
India carried birds to SE Asia from Madagascar (and by
extension from Africa) is not known. It certainly
transported ancient birds, but obvious descendants of these
may not have survived to modern times.
Africa was separated from western Laurasia by the
Tethys Sea until at least the Oligocene (Meulenkamp and
Sissingh 2003; Harzhauser et al. 2007; Allen and Arm-
strong 2008; Pook et al. 2009). A strong connection be-
tween African and SE Asian avifaunas is well known
(Olson 1973, 1979; Dinesen et al. 1994; Crowe et al. 2006;
Fuchs et al. 2006b; Jønsson et al. 2011; Ericson 2012;
Gonzalez et al. 2013), but it is difficult to determine how
and when bird groups were exchanged. Evidence from a
variety of plants and animals indicates that dispersal oc-
curred between Africa and SE Asia across Arabia and India
starting in the late Oligocene in what has been described by
Gonzalez (2012) as the ‘‘palaeotropical biotic exchange’’
(Morley 2000; Antoine et al. 2003; Zhou et al. 2012; Li
et al. 2013). Another ‘‘exchange’’ occurred at ap-
proximately 18–17 Ma during the height of the wet, warm
Miocene—this time by way of the Gomphotherium land-
bridge—and periodically thereafter (Barry et al. 1985,
1991; Rogl 1998, 1999; Koufos et al. 2005). The exchange
was closed toward the end of the Miocene with aridifica-
tion in the Middle East. Even during times when birds
could move back and forth between Africa and SE Asia,
the avifaunal connection was complicated by intervening
Laurasian and Indian avifaunas. Although Africa and SE
Asia share phasianids, cuckoos, trogons, hornbills, barbets,
woodpeckers, honeyguides, sub-oscines, among others,
these groups may well be Laurasian (Mayr 2005, 2014; but
see Ericson 2012). As such, they could have invaded Africa
and SE Asia independently. Thus, with the exception of a
few well-studied taxa, such as hornbills and oscine pas-
serines, it is difficult to stipulate with confidence a direct,
cross-India connection between Africa and SE Asia.
Numerous authors have also proposed that with respect to
some oscines, invasion may have occurred directly over the
Indian Ocean from Asia to Africa and vice versa (Jønsson
and Fjeldsa 2006; Samonds et al. 2012; Fjeldsa 2013;
Voelker et al. 2014).
In contrast, Australia’s influence on SE Asian birds is
much less speculative. At the beginning of the Cenozoic,
Australia was distant from SE Asia (Fig. 2), but by the end
of the Oligocene (approx. 23 Ma) the Australian plate had
pushed into the region of Sulawesi, closing the deep ocean
trench that had separated the two continents (Morley 2012;
Hall 2013). At this time, birds would have begun moving
from Australia to SE Asia (and vice versa) across inter-
vening islands (Barker et al. 2004; Schweizer et al. 2010;
Jønsson et al. 2011; Aggerbeck et al. 2014; Cibois et al.
2014), but the degree of avifaunal exchange is uncertain;
floral exchange at this point was minimal (Richardson et al.
2012). Extensive exchange between Australia and Sunda-
land would probably have proceeded with the Miocene,
Table 2 Number of resident land bird species shared between major
landmasses in Sundaland
Area Malay Peninsula Borneo Sumatra Java
Malay Peninsula –
Borneo 280 –
Sumatra 342 289 –
Java 205 174 222 –
Palawan 88 84 82 84
Based on the list of Sundaic species in the ESM 1 and the classifi-
cation of Gill and Donsker (2014)
J Ornithol
123
after fuller development of Wallacea (Fig. 3) caused by the
amalgamation of the Sundaic and Australian continental
plates (Stelbrink et al. 2012; Holt et al. 2013; de Bruyn
et al. 2014).
Geological history of Sundaland
Sundaland is a subregion of Malesia, which also includes
the Philippines, Wallacea, and New Guinea (van Steenis
1950; Whitmore 1981). The tectonic force that built
Malesia was the clash between the Pacific, Indian, Aus-
tralian, Asian, and many smaller continental plates, starting
at approximately 45 Ma. These events have been reviewed
by Robert Hall of the Royal Holloway University of
London (Hall 1998, 2002, 2009, 2012, 2013), and here we
rely heavily on his synopses. Plate subduction, twisting,
and collision resulted in the formation of modern Sumatra,
Java, parts of Borneo, Palawan, and thousands of conti-
nental and oceanic islands east of Sundaland. It also re-
sulted in the mingling of plants and animals of Asian and
Australian heritage across the entire region (Whitmore
1981, 1987; Morley 2012). To a large extent these geolo-
gical events coincided with the radiation of modern birds
and help explain the origin of the Sundaic avifauna. As
already noted, two especially important geological events
were the Eocene collision of the Indian and Asian plates
and the late Oligocene collision of the Australian and Asian
plates (Hall 2012; Morley 2012). The latter event cut the
connection between the Pacific and Indian oceans at the
Indonesian Throughflow, so that moisture previously car-
ried from the Pacific Warm Pool was shed on Sundaland
rather than further west in the Indian Ocean (Morley 2003,
2012; Hall 2013). The increase in rainfall accompanied
global warming in the Miocene, and it fed the rainforest of
Sundaland throughout most of the epoch. Toward the end
of the Miocene, the global climate became cooler and drier,
reaching a dry peak in the Pliocene. At the end of the
Pliocene, from 2.6 Ma onward, effects of global glacial
events came into play as powerful causal forces in avian
diversification in Sundaland.
Since the Mesozoic, the Sunda Core has extended as a
continental peninsula south from Asia (Fig. 3: 60 Ma). The
main central mountain range of the Malay Peninsula was in
place and above sea level throughout the Cenozoic. It and
other elevated parts of the Core, especially the Schwaner
Mountains of western Borneo and possibly areas in the
northern Java Sea, provided sediments to build surrounding
lowlands in the early Cenozoic. In contrast to this stable
center, the outer edges of Sundaland, namely, Sumatra,
Java, and eastern Borneo, experienced the effects of sub-
duction and mountain-building starting approximately
45 Ma and underwent much change.
Portions of western and northern Borneo were probably
emergent throughout the Cenozoic (Moss and Wilson
1998). At its beginning, Borneo was a fairly linear
promontory extending eastward from the Sunda Core
(Fig. 3). At approximately 20 Ma, mountain-building was
extensive on Borneo’s northern margin as the island rotated
counterclockwise on its axis. The growth rate of these
mountains was perhaps equivalent to that of the Himalayas
(Hall and Nichols 2002: 18), and the resulting erosion was
massive, filling large basins with sediment and creating the
northwestern, southeastern, and eastern lowlands of Bor-
neo. Subduction in the middle to late Miocene also created
a volcanic arc in the region of the Dent and Semporna
peninsulas, extending across the Sulu region to the
Philippines (Fig. 3: 10 Ma). The Meratus Mountains of
southeastern Borneo arose in the middle to late Miocene
(Witts et al. 2012). Mt. Kinabalu, Borneo’s most dominant
peak at 4095 m a.s.l., derives from granitic magma forced
into weak zones and fractures in Borneo’s northernmost
mountains. Subsequent uplift at approximately 7–8 Ma and
Fig. 2 Potential sources of birds in SE Asia (black arrows) at various
times in the Cenozoic: 60 Ma Laurasia, 40 Ma India, 30–10 Ma
Australia and Africa. For non-corvoid oscines (Passerides), biogeog-
raphers have speculated on a route to SE Asia first from Australia to
Africa over the Indian Ocean (approx. 40 Ma), and then more
recently from Africa to SE Asia (Ericson et al. 2003; Fuchs et al.
2006a; Jønsson and Fjeldsa 2006; Fjeldsa and Bowie 2008).
Continental positions were estimated from Li and Powell (2001) via
Schweizer et al. (2010)
J Ornithol
123
erosion of overlying softer rock (Choi 1996; Cottam et al.
2010) yielded the mountain we see today.
Sumatra experienced periods of mountain-building on
its southwestern side from the subduction of the Indian
plate, but the island also subsided and was largely inun-
dated several times. Volcanic activity was widespread in
the middle Eocene. From 30–25 Ma, Sumatra consisted of
mostly dry land, with small areas of high elevation and
some volcanoes, but from 20–10 Ma most of the lower
elevation sites were submerged, and only the mountains
remained above sea level. In the late Miocene, the Barisan
mountains rose and expanded (Fig. 3: 10 and 5 Ma). The
large islands off Sumatra’s southwestern coast (e.g., Nias
and Siberut) were probably connected to the main island
Fig. 3 Geography of Sundaland during the Cenozoic, based on maps provided by Hall (2013)
J Ornithol
123
since the middle Miocene, but this is less likely for islands
further to the east. At approximately 1 Ma, Sumatra ex-
perienced a large amount of volcanism along its western
side, and erosion of these mountains helped produce the
island’s current northern lowlands.
Java is a newer island than Borneo and Sumatra, having
emerged in its modern form only in the last 2 Ma, although
western Java existed as a land mass before then (Fig. 3).
The rocks underlying the modern island were roughly
aligned north–south with Sumatra at the beginning of the
Cenozoic, and its northern end (modern western end) was
connected and emergent at 30–25 Ma. From the end of the
Oligocene to about 5 Ma, however, Java was basically an
arc of volcanic islands rather than a single island. Like
Sumatra, it experienced a great deal of volcanism during
the last million years due to subduction. Unlike Sumatra,
however, it does not have a continuous mountain range; its
highlands and volcanos, which were previously separated
by water, are now separated by drier lowlands. Thus, the
mountains still act as geographic islands (Whitten et al.
1996).
Palawan’s history differs substantially from that of the
other Sunda islands. Palawan began as a continental frag-
ment that broke off southern Asia about 30 Ma and drifted
south until becoming lodged on the Sunda shelf about
10 Ma (Hall 1998, 2002; Zamoros and Matsuoka 2004;
Zamoros et al. 2008). Originally, Palawan was thought to
have been submerged during most of its journey, but
Blackburn et al. (2010) and Siler et al. (2012) have argued
on biological and geological grounds that parts of Palawan
were continuously subaerial, so that the island could have
acted as an ‘‘ark’’, transporting organisms from continental
Asia to Sundaland, Philippines, and Sulawesi. Hall’s most
recent maps (Fig. 3) depict Palawan as continuously
emergent as well. Much of modern-day Palawan was thrust
upward after it hit the Sunda shelf (Durkee 1993; Hall
2002; Yumul et al. 2009). Despite its attachment to the
Sunda shelf, Palawan may not have been connected di-
rectly by land to greater Borneo for at least 1 Ma, if ever
(Bintanja et al. 2005; Esselstyn et al. 2010).
In terms of avian evolution, the most important con-
clusion to be drawn from this geological summary is that
Borneo is by far the oldest and largest continuously sub-
aerial island in Sundaland. Its central mountain chain has
been dominant for at least 20 (perhaps 30) million years,
and its lowlands were extensive during that period. Su-
matra has come together as a stable, whole island only in
the last 5–10 Ma, and Java only in the last 2–5 Ma.
Geographic history of Sundaland
Geologists and paleontologists have made great strides in
identifying land areas and reconstructing climate and
habitats of Sundaland in the Cenozoic. Advances in un-
derstanding Sundaic geography have been reviewed and
updated for non-geologists by palynologist Robert J.
Morley of the Royal Holloway University of London
(Morley 1998a, b, 2000, 2011, 2012). Most of the evidence
for paleo climates and habitats comes from pollen derived
from drilling cores produced during hydrocarbon explo-
ration, but some also comes from lithographic indicators
such as coal, which was deposited during wet times, and
from paleosols (Morley and Morley 2013).
The Paleocene and Eocene of Sundaland were warm,
wet epochs, referred to as the Early Eocene Climatic Op-
timum (Yapp 2004). At this time, Sundaland would have
had extensive rainforest composed of a Laurasian flora
(Morley 2012). Modern plants did not occur in Sundaland
until the mid to late Eocene (Morley 2012), when many of
the older indigenous forms were replaced by invading In-
dian flora (Morley 2012: fig. 4.2). Forest birds of the early
and mid-Eocene would have been Laurasian as well, and
few would have been recognizable as modern taxa (Mayr
2014). Although cooling occurred toward the end of the
Eocene, it appears that the entire epoch would have suited
rainforest species, and we can expect they radiated sub-
stantially. From about 40 Ma, modern forms might have
included galliforms, cuckoos, owls, nightjars, frogmouths,
swifts, trogons, kingfishers, colies, coraciiforms, wood-
peckers, broadbills, and pittas (James 2005; Mayr 2005,
2014; Moyle et al. 2006a; Clarke et al. 2009; Ksepka and
Clarke 2009; Nesbitt et al. 2011).
During the Oligocene, Sundaland had seasonal forest in
its north and west and wetter forest along its eastern and
southern edges. Extensive drying in the mid-Oligocene
gave way to wetter forest in the equatorial zone and east
coast toward the end of the epoch (Morley 2012). Mid-
Oligocene (28 Ma) dry habitats are suggested by the ex-
tensive presence of conifers in the lowlands of the South
China Sea region and a lack of coal deposits (Morley 2012:
fig. 4.7). Sumatra also appears to have had a seasonal cli-
mate at this time. In the Java Sea, the early Oligocene was
wet but became drier as the epoch proceeded. Peatswamp
forest is evident in this region in the late Oligocene. Most
important for our story is that the mountains of equatorial
Borneo appear to have retained an ‘‘everwet’’ climate
throughout the epoch (Morley 2012).
At the end of the Oligocene, the collision between the
Australian and Asian plates closed the Indonesian
Throughflow, causing the warm, moist Pacific air to con-
centrate on Sundaland (Morley 2003). This moisture,
combined with global warming, ended the relatively cold,
dry Oligocene and began the long, warm, wet Miocene.
Extensive rainforest is thought to have developed across
Sundaland by the early Miocene, except perhaps in
southern Sumatra and western proto-Java (Morley 2012:
J Ornithol
123
fig. 4.7). Phylogenetic studies indicate that rainforest bird
groups, including galliforms, trogons, barbets, hornbills,
and oscine passerines, radiated substantially during this
epoch.
Sundaic bird biogeography from the Eocene to Pliocene
Here we describe patterns of Eocene–Pliocene evolution in
a few SE Asian rainforest bird groups that have been well
sampled and studied using modern phylogenetic methods.
One problem to be faced in reconstructing older biogeo-
graphic events in Sundaland, however, is that molecular
estimates of group ages (e.g., Brown et al. 2008; Wright
et al. 2008; van Tuinen 2009; Brown and Van Tuinen
2011) tend to be greater than those based on fossil or
geographic evidence of those ages (Feduccia 2003; Mayr
2013, 2014), sometimes by tens of millions of years. In SE
Asia, dating of phylogenetic events has depended on as-
sumed molecular rates because calibrating potentially more
appropriate rates is difficult given the region’s complex
geography and lack of rainforest bird fossils (Meijer 2014).
However, the timing of modern bird occurrence in Sun-
daland can be judged roughly by the age of modern
(crown) group fossils found elsewhere. Such fossils do not
occur in substantial numbers until the late Eocene (approx.
40 Ma; Mayr 2005, 2014). Also, newer molecular phylo-
genetic methods employing Bayesian relaxed clock ana-
lyses (Drummond et al. 2006; Drummond and Rambaut
2007; Ho and Phillips 2009) provide dates of bird-group
radiations that are more in accord with geological and
fossil evidence (Gonzalez 2012; Gonzalez et al. 2013; Stein
2013; Aggerbeck et al. 2014). These two lines of evidence
point circumstantially toward the Oligocene as the best
starting point for a discussion of the evolution of modern
Sundaic birds. At this time, the Eocene rainforest was re-
ceding because of a drier, cooler climate, and Eocene bird
groups probably faced substantial competition from Afri-
can and Australian invaders. After the Oligocene, with
climate amelioration, all phylogenetic studies agree that
modern rainforest bird groups flourished.
Galliformes Stein (2013) produced a comprehensive
phylogeny of galliforms employing[14,500 nucleotides of
mitochondrial and nuclear DNA isolated from 233 taxa,
with four fossils for calibration, and Bayesian relaxed clock
analyses. One of the most interesting early events evident
in Stein’s tree is the late Eocene–Oligocene connection
between the east African partridge Xenoperdix and the
Sundaic genera Caloperdix, Rollulus, and Arborophila
(Fig. 4). Such a relationship was predicted by Dinesen
et al. (1994) when they discovered Xenoperdix in the
mountains of Tanzania. Aging from the phylogeny com-
bined with modern distributions also suggests that
Caloperdix and Haematortyx (and possibly Rollulus) are
relicts that survived in Bornean refugia (Fig. 4). As with
other phylogenetic studies, Stein’s study demonstrates that
the late Miocene and Pliocene were most important to the
derivation of the modern pheasant and partridge species in
SE Asia.
Fruit pigeons and doves Sundaland’s main columbids are
fruit pigeons, among which Treron is the most speciose
group, being sister to all other fruit pigeons, doves, and
allies (including Chalcophaps, Turtur, Goura, Caloenas,
Trugon, Phapitreron, Hemiphaga, Lopholaimus, Gymno-
phaps, Ducula, and Ptilinopus), and appears to have arisen
in Asia (Johnson, personal communication). Ptilinopus
represents a Pacific radiation, with P. jambu reaching
Sundaland approximately 10 Ma (Cibois et al. 2014).
Parrots Parrots of Sundaland derived from Australia,
possibly in three waves toward the end of the Oligocene,
leading to Psittacula, Psittinus, and Loriculus. Psittacula
and Psittinus are relatively closely related to one another,
and Loriculus is related to lories (Schweizer et al. 2010).
Ground cuckoos Asian ground cuckoos (Carpococcyx)
form the sister group of couas (Coua) of Madagascar, and
the two genera diverged from one another 25–45 Ma
[Sorenson and Payne (2005) and personal communication].
Thus, it is tempting to say that the ancestors of Asian birds
rafted on India from Africa. However, cuckoos are possibly
a Laurasian group (Mayr 2005), and Madagascar is
Gondwanan. Thus, any number of evolutionary scenarios is
possible, including those of Carpococcyx and Coua being
relicts of a once widespread taxon or representative of in-
dependent invasions of Africa and Asia.
Trogons and barbets Trogons and barbets probably are
Eocene groups of Laurasia (Mayr 2005, 2014; but see
Ericson 2012). Both apparently invaded the Neotropics,
Africa, and Asian tropics at about the same time in the mid-
Cenozoic, thereby complicating the determination of basal
phylogenetic branching among the three tropical areas in
both groups (Moyle 2002, 2004, 2005; Johansson and
Erickson 2004). An early arrival of trogons and barbets in
SE Asia explains their parallel pattern of diversification in
the region. Both groups comprise what appears to be a
single relictual taxon—Apalharpactes in trogons and
Caloramphus in barbets—and a single speciose genus,
Harpactes and Megalaima (including Psilopogon), re-
spectively. Phylogenetic studies (Hosner et al. 2010; den
Tex and Leonard 2013) indicate a split between the re-
lictual and speciose genera approximately in the Oligo-
cene, and substantial diversification within Harpactes and
Megalaima in the Miocene (Fig. 5). One explanation for
this parallel pattern is that trogons and barbets proliferated
in the Eocene of SE Asia, then were largely extirpated in
J Ornithol
123
Fig. 4 Clades from the
galliform phylogeny of Stein
(2013) selected for their
Sundaic representation. Gray
boxes 95 % Highest posterior
densities of nodal ages. Where
taxa overlap, the branching
patterns here have been
corroborated by Wang et al.
(2013) and Sun et al. (2014)
Fig. 5 Similarity in pattern of
SE Asian trogon and barbet
phylogenies, given highly
speculative divergence dates.
Both groups are represented in
SE Asia by a singular, relictual
taxon of approximately
Oligocene age (Apalharpactes
in trogons and Caloramphus in
barbets), and both have
undergone substantial Sundaic
radiations in the Miocene and
Pliocene (Hosner et al. 2010;
den Tex and Leonard 2013)
J Ornithol
123
the colder, drier Oligocene, and finally re-radiated in the
Miocene. Apalharpactes and Caloramphus would be relicts
of the Eocene that survived the Oligocene (and Pliocene) in
rainforest refuges.
Hornbills Gonzalez et al. (2013) compared 1846 nu-
cleotides of the mitochondrial cytochrome b and nuclear
AK1 intron 5 to produce a dated phylogeny of all 61
species of hornbills. These authors calibrated the tree using
a combination of the coraciiform molecular rate (Pacheco
et al. 2011) and three hornbill-fossil dates. With the ex-
ception of Berenicornis (White-crowned Hornbill), they
found that Asian hornbills are monophyletic and invaded
from Africa in the late Oligocene. Berenicornis represents
a separate invasion from Africa in the early Miocene.
Gonzalez (2012) also estimated phylogenies of hornbill
food-trees to assess the co-evolution of the birds and their
food. The hornbills’ Asian invasion followed their devel-
opment of frugivory, and hornbills probably helped dis-
perse food-trees from Africa to SE Asia. An alternative
view had been proposed earlier by Viseshakul et al. (2011),
who believed that hornbills derived from Laurasia much
earlier (in the mid-Eocene) and dispersed Indian fruits to
insular SE Asia. The hornbills of modern Sundaland are
distributed in four main clades, with most members having
proliferated in the late Miocene.
Passerines Distribution, phylogeny, and the fossil record
suggest that Old World suboscines may have radiated in
the Eocene, only to be culled by the climate in the Oli-
gocene, and then largely replaced in the Miocene by
oscines. If true, the suboscines’ fate would mirror the
suspected replacement of other small-bodied Eocene
groups, such as colies and coraciiforms, by Miocene
oscines (Feduccia 1999; Mayr 2005; Clarke et al. 2009).
Evidence of Old World suboscines in the Eocene includes
phylogenetic estimates of lineage ages (Irestedt et al. 2001;
Moyle et al. 2006a) and fossils in the lower Oligocene of
Europe (Mayr 2013). Among Sundaic suboscines (broad-
bills and pittas), the frugivorous green broadbills (Calyp-
tomena) appear to be the only descendants of one ancient
lineage of broadbills, and the insectivorous broadbills
(Corydon and Eurylaimus) descendants of another. Pittas
are also ancient, but the modern taxa have diversified more
recently and extensively (or survived extinction more
successfully) than broadbills.
Oscines presumably arrived in SE Asia later than
suboscines, as indicated by phylogenetic and fossil
evidence (Ericson et al. 2003; Barker et al. 2004; Mayr
2005). Patterns of their likely derivation, dispersal, and
radiation have been reviewed by Fjeldsa (2013). Corvoid
oscines of SE Asia, i.e., members of Infraorder Corvides
(Cracraft 2014)—including orioles, vireos, whistlers,
minivets, cuckoo-shrikes, trillers, wood swallows, wood
shrikes, flycatcher-shrikes, Bornean Bristlehead, ioras,
fantails, drongos, monarchs, jays, and crows—most likely
reached Sundaland by island hopping from the Australian
region, starting in the late Oligocene, based on phyloge-
netic (e.g., Jønsson et al. 2008, 2010a, b, c, 2011; Fabre
et al. 2012; Aggerbeck et al. 2014) and geographic
evidence (Morley 2012; Stelbrink et al. 2012; Hall 2013).
Some of the corvoids then moved to Africa, probably
overland, and some of their descendants reinvaded SE
Asia.
The other SE Asian oscines are in the Infraorder
Passerides (Cracraft 2014) and include such groups as
parids, nuthatches, swallows, warblers, babblers, bulbuls,
flycatchers, starlings, flowerpeckers, and sunbirds. Their
route to SE Asia is more speculative than that of the
corvoids (Ericson et al. 2014). One proposal is that their
ancestors first invaded Africa from Australia in the Eocene
over the Indian Ocean, perhaps via the Kerguelen Plateau
or Indian Ocean islands (Fig. 2; Ericson et al. 2003; Fuchs
et al. 2006a; Jønsson and Fjeldsa 2006; Fjeldsa and Bowie
2008). Members of Passerides presumably arrived in SE
Asia overland after diversifying in Africa, although some
researchers have also suggested an invasion of SE Asia
from Africa directly over the Indian Ocean (Voelker et al.
2014). The earliest overland connection between Africa
and SE Asia would have been in the late Oligocene,
followed by periodic opportunities in the Miocene.
Molecular phylogenetic investigations have clarified the
relationships of the two iconic passerines, the Bornean
Bristlehead and Malaysian Rail-babbler, mentioned earlier
in this article. The Bristlehead is a classic corvoid, similar
in body plan (if not plumage) to Australian cracticids
(butcherbirds and magpies). It is probably a relict of the
Australian corvoid invasion of Sundaland, whose descen-
dants subsequently reached Africa (Ahlquist et al. 1984;
Moyle et al. 2006b; Jønsson et al. 2011; Aggerbeck et al.
2014). The Rail-babbler is sister to the African Chaetops
(rockjumpers), and together they are sister to the African
Picathartes (rockfowl) (Jønsson et al. 2007). Chaetops and
Picathartes are thought to be relicts of early Passerides
radiation in Africa (Ericson et al. 2014). Thus, the Rail-
babbler would be a relict of an early invasion of SE Asia by
ancestral African oscines.
Among Passerides that have proliferated in SE Asia, the
babblers (Timaliidae) are perhaps the best group for the
study of Sunda biogeography and community assembly
because of their taxonomic and morphological diversity,
approximately 275 species in 50 genera, and extensive
sympatry in rainforest habitats. Moyle et al. (2012)
provided a comprehensive phylogeny of the group and a
general biogeographic analysis. These authors found that
babblers are divided into four main clades—Leiothrichi-
nae, Timaliinae, Pellorneinae, and Zosteropidae—all of
J Ornithol
123
which presumably radiated in the Miocene (Fig. 6). As a
whole, Sundaic babblers appear to have a mainland Asian
origin. In Leiothrichinae, only a few taxa have invaded
Sundaland (some laughingthrushes and Brown Fulvetta,
Alcippe bruneicauda). The most diverse Sundaic groups,
Timaliinae and Pellorneinae, comprise clades that are
endemic and probably diversified in Sundaland (this is
especially true of the Pellorneinae), as well as taxa that
may have originated in Sundaland and invaded Indochina
and other mainland areas, such as the Scaly-crowned
Babbler (Malacopteron cinereum) and Grey-throated Bab-
bler (Stachyris nigriceps). The Zosteropidae differs from
other babblers in that most of the early diversification of
these birds may have been centered in Wallacea and the
Philippines (Cibois et al. 2002; Moyle et al. 2009). This
family also includes two Bornean endemics, Chlorocharis
and Oculocincta, which are aberrant morphologically but
close genetically to other white-eyes.
Among passerines arriving most recently in Sundaland
from Australia and the Pacific are Gerygone and Pachy-
cephala (Gardner et al. 2010; Jønsson et al. 2014). For the
most part, these invaders are coastal, but one species, P.
hypoxantha (Bornean Whistler), is endemic to the moun-
tains of Borneo.
Sundaland from the Pliocene to present
Regional endemism of Sundaic birds
During the peak of the Miocene, tropical forest reached
from the equator northeast as far as Japan and northwest as
far as India (Morley 1998b). The strong taxonomic con-
nection between Sundaic, Indochinese, and southern Chi-
nese avifaunas (such as shown in Fig. 1 of Packert et al.
2012) probably stems from this widespread Miocene
rainforest. Toward the end of the Miocene and especially in
the Pliocene, however, perhumid forest retracted with
global cooling and drying and became restricted to Sun-
daland (Morley 2012) and parts of northern Indochina (Li
and Walker 1986). Between these two areas, drier seasonal
forest developed. Within Sundaland during the Pliocene,
rainforest retreated still further, occurring mainly in Bor-
neo’s eastern mountains and its Makassar Strait and Ma-
hakam catchments on the eastern side of the island (Morley
and Morley 2011; Morley 2012; de Bruyn et al. 2014).
Rainforest persisted in eastern Borneo throughout the
Pliocene and Pleistocene, even when the rest of Sundaland
was drier. The forest situation in western Sundaland is not
as clear, but the mountains and islands of western Sumatra
were likely wet from orographic precipitation throughout
the Pliocene–Pleistocene. In Java, at approximately 2 Ma,
the eastern lowlands were savanna and nearly treeless, and
temperate flora was invading from the Australian side
(Morley 2012).
The separation of Sundaic and Indochinese rainforest for
the last 5–10 Ma (Morley 2012: Fig. 4.7) led to substantial
regional endemism (Table 1). Nevertheless, some taxa still
managed to move between Sundaland and Indochina
(Medway and Wells 1976; Hughes et al. 2003; Reddy
2008; Moyle et al. 2012), as evidenced by the low number
of endemic genera (23) in Sundaland (Table 3). Dispersal
of rainforest taxa between Sundaland and Indochina may
have occurred during relatively short wetter periods or
involved eurytopic intermediates able to move through
drier more open forest.
To reconstruct the development of Sundaic endemism, it
would be helpful to date vicariance and dispersal events
between Sundaland and Indochina, but few studies provide
the necessary information. Stein’s (2013) phylogeny of the
Galliformes (Fig. 4) has an approximate date for the di-
vergence of two relevant Arborophila species, A. javanica
(Java) and A. ardens (Hainan), which diverged ap-
proximately 5.1 Ma. Stein’s phylogeny also contains dates
relevant to intra-Sundaic diversification. Among the low-
land peacock–pheasants, Polyplectron schleiermacheri
(Borneo) diverged from P. malacense (Thai-Malay Penin-
sula) at approximately 4.4 Ma (see also Kimball et al.
2001), and among the lowland fireback pheasants, Lophura
Fig. 6 Outline of the babbler phylogeny (Moyle et al. 2012), with
Sundaic genera listed in major clades. Diversification times are highly
speculative
J Ornithol
123
bulweri (Borneo) diverged from its congeners (including
Sundaic and SE Asian taxa) approximately 7.4 Ma.
Two other studies that provide insight into the Sundaic–
Indochinese divergence are Reddy’s phylogeographic
comparisons of shrike-babblers and scimitar babblers
(Reddy 2008; Reddy and Moyle 2011). Among scimitar
babblers, she found that the endemic Sundaic species Po-
matorhinus montanus (Borneo, Sumatra, Java, and the
Malay Peninsula) was isolated near the beginning of the
Pleistocene from its Indochinese sister group, which con-
tains all other Pomatorhinus species. Among shrike-bab-
blers, she discovered two rounds of diversification. The
Javan species Pteruthius aenobarbus and P. flaviscapis
diverged from their closest Indochinese relatives at 4 and
2.5 Ma, respectively.
Finally, first Johansson et al. (2007) and then Packert
et al. (2012) examined the relationships of Sundaland’s
Phylloscopus and Seicercus warblers as tangents to their
Himalayan studies. As with shrike-babblers and scimitar
babblers, Sundaland has few representatives of these
otherwise speciose groups. P. trivirgatus (Mountain
Leaf-Warbler) diverged from its closest mainland rela-
tives between the late Pliocene and the mid-Pleistocene.
The Sundaic yellow-breasted warblers (S. montis, S.
castaniceps, and S. grammiceps) diverged as a group in
the late Miocene to Pliocene, but S. castaniceps has
maintained or regained a wide SE Asian distribution.
Opportunities to date Indochinese–Sundaic vicariance
and dispersal will accrue with more studies. Nevertheless,
doubts about the accuracy of molecular dating without
fossils or geographic evidence for calibration, and the in-
herent vagility of birds, will always plague efforts at
quantification.
The advent of high-latitude glaciation
Starting in the late Pliocene (3.2–2.6 Ma) and continuing
throughout the Quaternary, dramatic increases in the am-
plitude of climatic oscillations caused periods of global
cooling that resulted in long-lasting, high-latitude glacial
events (approx. 40,000–100,000 years each) separated by
relatively short interglacials (approx. 10,000–15,000 years
each), such as the one we are currently experiencing
(Kashiwaya et al. 2001; Zachos et al. 2001; deMenocal
2004; Bintanja et al. 2005). For Sundaland, the conse-
quences of these events were dramatic: lower sea levels as
arctic glaciers tied up oceanic water, a substantially in-
creased lowland area as the Sunda shelf emerged subaeri-
ally, connection of islands with one another and the
mainland, decreased oceanic influence on the interior areas
of Sundaland as the subcontinent enlarged, and increased
areas of montane habitat as mountain forests descended in
elevation with colder temperatures (Hanebuth et al. 2000;
Voris 2000; Wilson et al. 2000; Bird et al. 2005; Cannon
et al. 2009). During interglacials, sea level rose as arctic
glaciers melted, and the Sunda islands became isolated by
ocean waters, as they are today. The overall effect of these
alternating events was not only to isolate and, conversely,
join islands, but to shift the position, size, and types of
habitats covering Sundaland, creating diverse scenarios of
organismal vicariance and colonization.
The biogeography of Sundaland with respect to
Quaternary sea-level changes has been well reviewed
(Whitmore 1981; Heaney 1986; Whitmore 1987; Voris
2000; Woodruff and Turner 2009; Cranbrook 2010; Loh-
man et al. 2011; Morley 2012; de Bruyn et al. 2014;
Wurster and Bird 2014) and is only briefly covered here.
One issue, however, requires special emphasis because it
has been controversial and bears importantly on the evo-
lution of Sundaic birds: whether a drier, open habitat—
originally described as a ‘‘savanna corridor’’ (Heaney
1991; Bird et al. 2005)—occupied the interior of Sundaland
during periods of glacial perturbation. Existence of a dry
Sundaic interior is supported by some lines of evidence, but
disputed by others.
Our molecular phylogeographic studies of rainforest
birds reveal patterns of vicariance and dispersal that are
consistent with alternating periods of drier and wetter
Table 3 Endemic bird genera
of Sundaland
Based on the list of Sundaic
species in Electronic
Supplementary Material 1 and
the classification of Gill and
Donsker (2014)a B, Borneo; J Java; S, Sumatra;
M, Malay Peninsulab Paraphyletic genera (Moyle
et al. 2009; den Tex and
Leonard 2013)
Genus Areaa
Melanoperdix S M B
Rhizothera S M B
Haematortyx B
Hydrochous J S M B
Apalharpactes J S
Rhinoplax S M B
Berenicornis S M B
Reinwardtipicus J S M B
Psilopogonb S M
Caloramphus S M B
Pityriasis B
Urosphena B
Oculocinctab B
Chlorocharisb B
Chlamydochaera B
Platylophus J S M B
Platysmurus S M B
Eupetes S M B
Setornis S B
Tricholestes S M B
Kenopia S M B
Psaltria J
Leucopsar J
J Ornithol
123
forests in the interior of Sundaland in the Pleistocene. The
area of Sundaland between Borneo, Sumatra, and the
Malay Peninsula was occupied by a dry habitat in the early
to mid-Pleistocene which was sufficiently large to force
rainforest birds into refugia in eastern and western Sun-
daland. During other times (mid-Pleistocene onward?),
drying of Sundaland’s interior was not pervasive enough to
keep rainforest birds in refugia.
Few biogeographers have made the distinction between
the effects of different glacial events when discussing the
‘‘savanna corridor’’ because most arguments have focused
on the Last Glacial Maximum (LGM; approx. 18–21
thousand years ago). Indeed, failure to distinguish between
the effects of different glacial events seems to be the main
cause of disagreement. Only recently has this source of
confusion been emphasized. Morley (2012), and more re-
cently de Bruyn et al. (2014), appreciated the need to
consider the complexity of multiple glacial events, of
which about ten occurred in the last 1 million years and
about 20 (less severe) in the preceding 1 million years
(Augustin et al. 2004; deMenocal 2004). These events
varied in force and, thus, in their effects on habitats.
Sundaland’s forest habitats during glacial events
The data used to identify terrestrial Pleistocene habitats
derive from the fields of geology, palynology, paleon-
tology, and phylogeography. The first three sources usually
provide fairly accurate information on specific sites and
times. Thus, we know, for example, that parts of Sundaland
had a dry, open habitat at certain times and places, such as
in central to eastern Java in the late Pliocene through the
mid-Pleistocene (Cranbrook 2000; van den Bergh et al.
2001; Bettis et al. 2009; Louys and Meijaard 2010; Morley
2012) and on the Malay Peninsula during the LGM
(Wurster et al. 2010). We also know that parts of Sumatra
had wet, closed forest in the late Pleistocene (Louys and
Meijaard 2010). However, these data generally do not bear
on the most important areas for dispersal of rainforest birds
among the main islands, i.e., habitats intervening between
Sumatra, Borneo, and the Malay Peninsula.
The existence of a dry interior in Sundaland is supported
by mid-Pleistocene fossils of the Javan megafauna, includ-
ing extinct proboscideans, rhinos, cattle, buffalo, hog deer,
antelope, and hyenas. A dry habitat in central Sundaland was
required for some of these animals to reach Java from Asia in
the late Pliocene to early Pleistocene (Medway 1972; Hea-
ney 1991; Cranbrook 2000, 2010; Morley 2000; van den
Bergh et al. 2001; Meijaard 2003). Additional circumstantial
evidence is supplied by phylogeographic and distributional
studies of populations of rainforest animals and plants. In
these cases, rainforest populations were clearly divided be-
tween eastern and western Sundaland, apparently by
inhospitable (presumably dry, open) habitat in central Sun-
daland, and this subdivision almost always has been at-
tributed to the early Pleistocene, such as for mammals (Ruedi
1996; Zhi et al. 1996; Brandon-Jones 1998; Fernando et al.
2003; Gorog et al. 2004; Meijaard and Groves 2004; Hirai
et al. 2005; Steiper 2006; Patou et al. 2010; Wilting et al.
2012; Wurster and Bird 2014), reptiles and amphibians
(Wilting et al. 2012), fish (Ryan and Esa 2006), ants (Quek
et al. 2007), termites (Gathorne-Hardy et al. 2002), and trees
(Banfer et al. 2006; Iwanaga et al. 2012; Ohtani et al. 2013).
In contrast to megafaunal and phylogeographic data, only
a few studies dispute the existence of an extensive dry habitat
in central Sundaland. Paleo-habitat modeling and some
geological and organism–distribution studies (Hu et al.
2003; Kershaw et al. 2007; Cannon et al. 2009; Slik et al.
2009; Prentice et al. 2011; Handiani et al. 2013) have found
evidence of substantial rainforest (or at least wet habitats)
across Sundaland in the LGM. Other biogeographers have
presented nuanced perspectives of paleo-environmental
distributions based on thorough reviews of the literature and
sometimes quantitative examinations of fossil data (Mei-
jaard and van der Zon 2003; Louys and Meijaard 2010). Such
studies have uncovered conflicting evidence, especially on
Borneo (e.g., at Niah), where numerous and varying types of
paleo-data are available, and the Malay Peninsula, where
paleo-sites are few and dating uncertain. Such studies indi-
cate the possibility of a mosaic of habitats (in the LGM, but
presumably also during earlier glacial events), which may
have supported movement or isolation of rainforest and dry-
habitat taxa differentially across Sundaland at various times
(Meijaard and van der Zon 2003).
Regardless of the uncertainty about the nature and
timing of habitats at circumscribed sites and times, the
overwhelming weight of evidence indicates a fairly
straightforward outline of Pleistocene habitat geography
(Fig. 7). Late in the Pliocene or early in the Pleistocene,
drier, open forest or savanna must have existed in Sunda-
land, at one or multiple times. The drier habitat provided
the means for elements of the Asian megafauna to reach
Java, and it subdivided Sundaic rainforest into refugia in
eastern and western Sundaland. These refugia most likely
were associated with mountains, coastal areas, and islands,
where the effects of orographic rainfall would be the
greatest (Newsome and Flenley 1988; Stuijts et al. 1988;
Stuijts 1993; Brandon-Jones 1996, 1998; Morley 2000;
Gathorne-Hardy et al. 2002; Gorog et al. 2004; Wilting
et al. 2012). Lower elevation sites associated with moun-
tains would also have benefitted from direct rainfall or
indirect watering from montane runoff. We know from
palynological data that a rainforest refuge existed in eastern
Borneo from the late Miocene to the present (Morley and
Morley 2011; Morley 2012). More recently—almost cer-
tainly in the LGM, when modeling shows extensive
J Ornithol
123
rainforest across equatorial Sundaland (Cannon et al.
2009), as well as during other glacial events of the last
1 Ma, when the Sundaic interior likely comprised large
sections of kerangas and kerapah (everwet) forests (Morley
2012)—the savanna populations in Java were cut off from
Indochina by perhumid forests, and some refugial rain-
forest populations in eastern and western equatorial Sun-
daland were able to come together.
In the following section, we describe how this rough
approximation of events and timing applies to the birds we
have studied. However, we add the caveat that the differ-
ential effects of glacial events, the likelihood of a mosaic of
habitats, the uncertainty of molecular dating, and the id-
iosyncratic biology of individual bird taxa make our sce-
nario preliminary and almost certainly not universal.
Periodic land and habitat bridges may have allowed some
bird species—but not others—to move from one island to
another. Subsequent glacial events may or may not have
allowed sufficient dispersal to homogenize gene pools.
Small founding sizes of populations, low gene flow, and
unpredictable selection (e.g., sexual selection) would also
have helped drive diversification.
Phylogeography of Sundaic birds
Bornean lowland taxa
Sabah, the Malaysian state in northeast Borneo (Fig. 7),
has an unusually large number of endemic lowland birds
relative to the rest of Borneo, despite continuous rainforest
across the entire island (Sheldon et al. 2001, 2009a; Mann
2008). To investigate the cause of the marked lowland
parapatry in birds, we have conducted molecular phylo-
geographic studies on about 20 passerine species (Moyle
et al. 2005, 2011; Sheldon et al. 2009b; Lim et al. 2010,
2011, 2014; Oliveros and Moyle 2010; Lim and Sheldon
2011; Gawin 2014; Chua et al. 2015). These studies sug-
gest that many of Borneo’s lowland populations were
subdivided early in the Pleistocene into rainforest refugia,
probably when the interior of the Sundaland was drier
(reviewed in Gawin et al. 2014). Subsequently, probably in
the mid-Pleistocene and certainly during more recent gla-
cial events when the interior of Sundaland was wetter
(Cannon et al. 2009), refugial rainforest populations came
together to form a contact zone near the Sabah border
(Fig. 7). This vicariance–dispersal dynamic is illustrated by
the shama species group comprising Copsychus mal-
abaricus and C. stricklandii (Mees 1986, 1996; Lim et al.
2010, 2011, 2014; Gawin 2014; Chua et al. 2015).
Copsychus malabaricus has a black crown. It occurs
from India and southern China to Java, and it inhabits most
of Borneo. C. stricklandii has a white crown and is divided
into two subspecies: C. s. stricklandii in NE Borneo and C.
s. barbouri on Maratua, a small oceanic island ap-
proximately 50 km off Borneo’s east coast (Fig. 7). C.
stricklandii differs from C. malabaricus by a mitochondrial
ND2 gene sequence distance of [3 %. C. stricklandii’s
subspecies differ from each other by an ND2 distance of
2 %. The presence of a genetically divergent white-
crowned population on Maratua Island suggests that in the
early Pleistocene C. stricklandii occurred on Borneo adja-
cent to Maratua, i.e., it provided the stock for the invasion of
Maratua. Hence in Fig. 7, we have located the refugial C.
stricklandii population (in yellow) on Borneo’s east coast
adjacent to Maratua. Rainforest in this area is supported by
palynological and other data (Gathorne-Hardy et al. 2002;
Morley and Morley 2011). Subsequently, C. stricklandii on
mainland Borneo moved or retreated to the region of Sabah.
Its movement would have been in response to a habitat
change because many other Bornean endemics also cur-
rently occupy Sabah. Meanwhile, C. malabaricus must
have inhabited an allopatric refuge in western Sundaland.
This refuge could have been in any of several perhumid
sites in mountains or coastal areas (a few possibilities are
indicated in Fig. 7). Based on genetic similarity among C.
malabaricus populations from Sumatra, the Malay Penin-
sula, and western Borneo, this species must have moved
freely in western Sundaland in the mid to late Pleistocene,
approximately 0.5–1.0 Ma (Lim et al. 2010, 2011). In due
course, probably quite recently, C. malabaricus moved
across Borneo to meet and hybridize (to a limited degree)
with C. stricklandii near the Sabah border (Davison 1999;
Collar 2004; Gawin 2014).
Fig. 7 Map of Sundaland showing the approximate current distribu-
tion of endemic northeastern Bornean bird populations (orange) and
locations of possible Pleistocene rainforest refuges (yellow), including
Maratua Island off eastern Borneo (circle). The phylogeography of
Copsychus, highlighted in the text, concerns C. stricklandii, currently
occurring in the orange region of Borneo and on Maratua Island, and
C. malabaricus, currently occurring throughout the rest of Sundaland
(except Palawan) and across southern mainland Asia
J Ornithol
123
A similar dynamic occurred in Copsychus saularis, the
Oriental Magpie-Robin (Mees 1986, 1996; Sheldon et al.
2009b; Lim et al. 2010; Gawin 2014). Populations of this
species were also apparently subdivided in the early
Pleistocene. The main difference between magpie-robins
and shamas is that populations of magpie-robins from
western Borneo, Sumatra, and Java have more recently and
thoroughly intermixed, based on their genetic distances and
plumage morphology. Populations of eastern and western
Bornean magpie-robins have also experienced much
greater hybridization in their contact zone than have the
shamas (Gawin 2014), presumably because—as birds of
forest edge—they traversed Borneo more quickly than the
closed forest shamas. Analogous population interactions
also occurred in both Copsychus species on Java (Mees
1986, 1996; Gawin 2014).
To reiterate, the distribution and population genetics of
these Copsychus species strongly suggest vicariance in the
early Pleistocene, presumably by the inhospitable habitat.
More recently, these populations have come together be-
cause of continuous perhumid forest intervening in central
Sundaland (Cannon et al. 2009; de Bruyn et al. 2014:
Fig. 2).
Bornean montane taxa
The montane avifauna of Sundaland presents a more baf-
fling array of biogeographic patterns than the lowland
avifauna, making it difficult to reach general conclusions
about its evolutionary history. On Borneo, montane taxa
include widespread species (e.g., Turdus poliocephalus),
Himalayan taxa (Garrulax, Yuhina, Seicercus, Phyllosco-
pus, and Pycnonotus flavescens), ancient monotypic genera
(e.g., Haematortyx and Caloperdix), younger monotypic
genera (e.g., Chlamydochaera, Oculocincta, and Chlor-
ocharis), and many endemic representatives of Sundaic or
widespread SE Asian groups (e.g., Rhizothera dulitensis,
Arborophila hyperythra, Spilornis kinabaluensis, Collo-
calia dodgei, Harpactes whiteheadi, Megalaima eximia, M.
monticola, M. pulcherrima, Calyptomena hosii, Calyp-
tomena whiteheadi, Pitta arquata, etc.). Enigmatic para-
patric distributions also occur between montane taxa and
their lowland congeners. For example, the lowland spi-
derhunter Arachnothera modesta is replaced in the moun-
tains of western Borneo by its close congener, the endemic
A. everetti. In Sabah, however, A. modesta is absent and A.
everetti occupies both the lowlands and mountains (Shel-
don et al. 2001; Moyle et al. 2011). A similar pattern oc-
curs in the leafbirds Chloropsis cochinchinensis and C.
kinabaluensis, but with a twist (Wells et al. 2003; Sheldon
et al. 2009a; Moltesen et al. 2012). C. cochinchinensis
occurs in the lowlands of western Borneo and is replaced in
the western mountains by its close congener, the endemic
C. kinabaluensis. As with the lowland A. modesta, the
lowland C. cochinchinensis is absent in Sabah, but unlike
the ubiquitous Sabahan A. everetti, C. kinabaluensis re-
mains restricted to the mountains of Sabah and is not found
in the lowlands.
Despite its idiosyncrasies, the montane avifauna of Bor-
neo does exhibit a few predictable biogeographic patterns.
One such pattern is that Bornean montane species have
arisen primarily in allopatry, not in parapatry (i.e., not
through ecological speciation on an elevational gradient).
Although elevational parapatry of congeners is common on
Borneo (e.g., between the lowland swiftlet Collocalia es-
culenta and the montane endemic C. dodgei), the parapatric
congeners are not sister taxa. Sister taxa of Bornean montane
endemics have always been found to occur on other islands:
for example, Harpactes whiteheadi (Whitehead’s Trogon)
and H. ardens (Philippine Trogon), Arachnothera juliae
(Whitehead’s Spiderhunter) and A. clarae (Naked Faced
Spiderhunter) of the Philippines; A. everetti (Bornean Spi-
derhunter) and A. affinis (Streaky-breasted Spiderhunter) of
Java; Collocalia dodgei (Bornean Swiftlet) and C. lynchi
(Cave Swiftlet) of Java; Enicurus borneensis (Bornean
Forktail) and E. leschenaulti (White-crowned Forktail) of
Java; Chlamydochaera jefferyi (Fruithunter) and Cochoa
(cochoas) of Sumatra and Java; Chlorocharis emiliae
(Mountain Black-eye) and Zosterops montanus (Mountain
White-eye) of the Philippines, Wallacea, and other Greater
Sunda Islands (Klicka et al. 2005; Moyle et al. 2005, 2008,
2009, 2011; Hosner et al. 2010). When closely related con-
geners are elevationally parapatric, the lowland species ap-
pears to be a recent invader that has restricted the montane
species to higher elevation through competition (although
competition has not been directly demonstrated). This sce-
nario is highly reminiscent of the predictions of taxon cy-
cling (Ricklefs and Cox 1978).
The best studied Bornean montane endemic in terms of
phylogeography is Chlorocharis emiliae, the Mountain
Black-eye (Gawin et al. 2014). This species is a typical
white-eye (Zosteropidae), most closely related to Z. mon-
tanus (Moyle et al. 2009). It has a sky island distribution,
occurring on Borneo’s highest peaks in the central moun-
tain chain, but also on top of two outlying mountains in
western Borneo (Fig. 8). Molecular and morphological
comparisons of Chlorocharis populations indicate that it is
subdivided in the same way (including timing) as many
lowland taxa of Borneo, i.e., the Sabah population is dis-
tinct from that inhabiting the rest of the island (Ramji et al.
2012; Gawin et al. 2014). The pattern and timing of sub-
division suggest that the same Pleistocene forces that cre-
ated lowland subdivision and parapatry on Borneo—
vicariance during dry times and subsequent dispersal dur-
ing wetter times—also created montane subdivision and
parapatry on the island. Some other montane species that
J Ornithol
123
are similarly subdivided (based on their subspecific tax-
onomy) are: Garrulax treacheri (Chestnut-hooded Laugh-
ingthrush), Alophoixus ochraceus (Grey-cheeked Bulbul),
and Stachyris nigriceps (Grey-throated Babbler).
Given that orographic precipitation is thought to have
watered lowland rainforest associated with mountains
during dry periods of the Pleistocene, it makes sense that
the wet mountains themselves would have acted as refuges
for montane species. The difficulty then is determining
which mountains served as refuges for subdivided Bornean
montane bird populations. We know from pollen history
that the northeastern mountains (basically in Sabah) of-
fered refuge. The other(s) may have been in one or more
sites across the island (Fig. 7): the Pueh Range, Schwaner
Range (Gorog et al. 2004), or Meratus Range (Gathorne-
Hardy et al. 2002). The Meratus Range seems most likely
from an orographic rainfall standpoint (near the coast,
away from central Sundaland), but in terms of the current
phylogeographic structure of birds, the western or central
mountains of Borneo present a more attractive alternative.
Molecular comparisons of birds from the Meratus Range
should shed light on this issue.
Javan taxa
Because of its young age, Java is home to relatively recent
invaders from Sumatra, Borneo, and Wallacea. Most
interestingly for the reconstruction of Sundaic biogeography,
some of these recent invaders are more closely related to
species in Indochina than to those on other Sunda islands.
Disjunct Javan and Indochinese species include mainly dry-
habitat birds, such as: Dendrocopus analis (Freckle-breasted
Woodpecker), Psittacula alexandri (Red-breasted Parakeet),
Pericrocotus cinnamomeus (Small Minivet), Prinia polychroa
(Brown Prinia), Orthotomus sutorius (Common Tailorbird),
Timalia pileata (Chestnut-capped Babbler); however, some
rainforest species are disjunct as well, such as Tesia
cyaniventer (Grey-bellied Tesia). There are also some closely
related species pairs: Locustella montis (Javan Bush Warbler)
and L. mandelli (Russet Bush Warbler), among others. Only a
few studies have compared these disjunct populations using
molecular methods, e.g., Psittacula alexandri (Kundu et al.
2012), Pericrocotus cinnamomeus (Jønsson et al. 2010c),
Orthotomus sutorius (Sheldon et al. 2012), and Locustella
montis (Alstrom et al. 2011). In each case a close phylogeo-
graphic relationship is evident. In our study of tailorbirds, for
example, we found an uncorrected ND2 sequence divergence
of only 0.8–1.0 % between individuals of Orthotomus sutorius
of Java and Singapore, to which this tailorbird has recently
invaded from the north due to deforestation and drying of the
Malay Peninsula (Medway and Wells 1976). This genetic
distance suggests (but by no means proves) that the tailorbird’s
disjunction occurred in the mid-Pleistocene. As noted earlier,
Reddy (2008) discovered older Javan–Indochinese connec-
tions in shrike-babblers (early Pleistocene and Pliocene).
The ancestors of the disjunct taxa most likely reached
Java across the interior of Sundaland. Rainforest birds, like
Tesia, would also have invaded Sumatra and Borneo and
subsequently been extirpated (perhaps through competi-
tion) on those islands. Dry habitat birds would have
reached Java in the same manner as the Javan megafauna,
through dry interior parts of the Sunda shelf, starting per-
haps in the Pliocene. Judging from the taxic variety (and
presumed genetic variation) between pairs of dryland dis-
juncts, invasion may have occurred in waves, with those
disjunct at the population level arriving most recently. As
with the megafauna, Java’s dry-habitat birds would have
been cut-off from their sister taxa in Indochina during in-
terglacials and during low sea-level events when Sunda-
land’s interior forest was perhumid (Cannon et al. 2009;
Morley 2012). Further molecular comparisons between
Javan and Indochinese populations should yield a rich crop
of biogeographic discovery, especially in dating invasions
and establishing the timing of habitat occurrence.
Conclusions
A major conclusion of this review is that Borneo played the
preeminent role in the evolution of rainforest birds in
Fig. 8 The sky island distribution of Mountain Black-eye (Chlor-
ocharis emiliae) on Borneo. Triangles Known populations, white
circles populations in the two main clades, which are separated by a
mitochondrial ND2 sequence difference of 2.5 %. The northernmost
clade, in Sabah, likely represents the influence of an early Pleistocene
refuge in the vicinity of Mt. Kinabalu. The southwestern clade’s
Pleistocene refuge is unknown, but could be in any site subject to
orographic rainfall during dry periods
J Ornithol
123
Sundaland (de Bruyn et al. 2014). Borneo’s evolutionary
power stems from the long-term existence of its eastern
mountains and their adjacent lowlands, which served as
refuges during the drier, cooler Oligocene, Pliocene, and
Pleistocene and helped foment taxa from the Miocene
through Pleistocene. The island’s refuges probably pre-
served several relicts, including the montane Haematoryx
sanguiniceps (Crimson-headed Partridge) and Caloperdix
oculeus (Ferruginous Partridge), and possibly the lowland
Rollulus roulroul (Crested Partridge) and Calorhamphus
(brown barbets). Bornean mountains probably also were
the center of Miocene bird evolution in Sundaland. In the
Miocene, lowland rainforest extended widely in southern
Asia, but Borneo’s mountains were isolated from those at
higher latitude throughout the epoch (Fig. 3). In contrast to
Borneo, Sumatra or Java did not have substantial land area
until 10 Ma or 5 Ma, respectively, when the wet warmth of
the Miocene was waning. The role of eastern Java, arising
after 5 Ma, would have been very small in terms of the
proliferation of major Sundaic groups, but eastern Java
would have been important in preserving disjunct In-
dochinese taxa.
The Pleistocene brought a different dynamic to Sunda-
land. Starting early in the Pleistocene and continuing
probably until at least 800 ka, glacial events caused enough
habitat drying in central Sundaland to subdivide rainforest
into refugia and to allow dry-habitat birds and mammals to
reach Java from Indochina. Subsequently, glacial events
featured enough perhumid habitat to connect previously
vicariated rainforest populations, creating the parapatry we
see in both the lowlands (e.g., on Borneo, between Sabah
and elsewhere) and mountains (e.g., on Borneo, in Collo-
calia, Enicurus, Chloropsis, Arachnothera, etc.). The dy-
namics between a drier and wetter Sundaic interior was
probably not simply one period of separation and one pe-
riod of connection, but rather a complex interplay of iso-
lation on islands (including islands of habitat) and
colonization events via land or habitat bridges. Also at play
in diversification would have been the influence of
relatively small population sizes, low gene flow, and be-
havioral idiosyncrasies of each species.
Future research needs
Directions of future research are clear. At the population
level, to reconstruct the evolution of Sundaic birds effec-
tively we need extensive sampling from areas for which
there are almost no modern bird specimens: Kalimantan in
southern Borneo, Java, Sumatra, the Malay Peninsula, and
Thailand. For population studies, we also need to move
from Sanger sequencing of mitochondrial DNA and slowly
evolving nuclear genes to next-generation sequencing
methods. The latter will provide more independent nuclear
loci for comparison and, consequently, more precise esti-
mates of phylogeographic parameters, such as population
size, gene flow, and divergence time. For phylogenetic
studies, we need more extensive taxonomic sampling,
which again requires more and better collections. Finally,
we need more information on paleo-habitats that occurred
during early Pleistocene glacial events. Such data will most
likely be provided by increased and improved modeling
methods and continued work on fossil plants and pollen.
Acknowledgments We especially thank Franz Barlein, Erik
Matthysen, David Winkler, and the International Ornithological
Congress for inviting FHS to present a plenary lecture and write this
paper. We also thank Robert Hall for providing his Cenozoic maps
(Fig. 3) and for giving permission to use them, and JC Gonzalez and
Will Stein for sending material from their PhD theses. We are ex-
tremely grateful for input from Ed Braun, Clare Brown, Ryan Burner,
Vivien Chua, Geoff Davison, Dency Gawin, Jake Esselstyn, Kevin
Johnson, Rebecca Kimball, John Mittermeier, Quentin Phillipps, Rick
Prum, Mustafa Abdul Rahman, Mohamad Fizl Sidq Ramji, Roselyn
Remsen, Van Remsen, Frank Rheindt, Subir Shakya, Mike Sorenson,
Katie Stryjewski, and David Wells. This project has been supported
by the Malaysian Chief Minister’s Department and numerous gov-
ernment departments in Sabah and Sarawak. Funding was provided
by NSF DEB-0228688, NSF DEB-1241059, Coypu Foundation of
Louisiana, National Geographic Society, Louisiana State University,
American Museum of Natural History, University of Kansas, Sabah
Parks, Sabah Museum, and the Universiti Malaysia Sarawak.
References
Aggerbeck M, Fjeldsa J, Christidis L, Fabre PH, Jønsson KA (2014)
Resolving deep lineage divergences in core corvoid passerine
birds supports a proto-Papuan island origin. Mol Phylogenet
Evol 70:272–285
Ahlquist JE, Sheldon FH, Sibley CG (1984) The relationships of the
Bornean Bristlehead (Pityriasis gymnocephala) and the Black-
collared Thrush (Chlamydochaera jefferyi). J Ornithol
125:129–140
Aitchison JC, Ali JR, Davis AM (2007) When and where did India
and Asia collide? J Geophys Res 112:B05423
Allen MB, Armstrong HA (2008) Arabia-Eurasia collision and the
forcing of mid-Cenozoic global cooling. Palaeogeogr Palaeocli-
matol Palaeoecol 265:52–58
Alstrom P, Fregin S, Norman JA, Ericson PG, Christidis L, Olsson U
(2011) Multilocus analysis of a taxonomically densely sampled
dataset reveal extensive non-monophyly in the avian family
Locustellidae. Mol Phylogenet Evol 58:513–526
Antoine P-O, Welcomme J-L, Marivaux L, Baloch I, Benammi M,
Tassy P (2003) First record of Paleogene Elephantoidea
(Mammalia, Proboscidea) from the Bugti Hills of Pakistan.
J Vertebr Paleontol 23:977–980
Augustin L, Barbante C, Barnes PRF, Barnola JM, Bigler M,
Castellano E, Cattani O, Chappellaz J, DahlJensen D, Delmonte
B, Dreyfus G, Durand G, Falourd S, Fischer H, Fluckiger J,
Hansson ME, Huybrechts P, Jugie R, Johnsen SJ, Jouzel J,
Kaufmann P, Kipfstuhl J, Lambert F, Lipenkov VY, Littot GVC,
Longinelli A, Lorrain R, Maggi V, Masson-Delmotte V, Miller
H, Mulvaney R, Oerlemans J, Oerter H, Orombelli G, Parrenin F,
J Ornithol
123
Peel DA, Petit JR, Raynaud D, Ritz C, Ruth U, Schwander J,
Siegenthaler U, Souchez R, Stauffer B, Steffensen JP, Stenni B,
Stocker TF, Tabacco IE, Udisti R, van de Wal RSW, van den
Broeke M, Weiss J, Wilhelms F, Winther JG, Wolff EW,
Zucchelli M, Members EC (2004) Eight glacial cycles from an
Antarctic ice core. Nature 429:623–628
Banfer G, Moog U, Fiala B, Mohamed M, Weising K, Blattner FR
(2006) A chloroplast genealogy of myrmecophytic Macaranga
species (Euphorbiaceae) in Southeast Asia reveals hybridization,
vicariance and long-distance dispersals. Mol Ecol 15:4409–4424
Barker FK, Cibois A, Schikler P, Feinstein J, Cracraft J (2004)
Phylogeny and diversification of the largest avian radiation. Proc
Natl Acad Sci USA 101:11040–11045
Barry JC, Johnson NM, Raza SM, Jacobs LL (1985) Neogene
mammalian faunal change in southern Asia: correlations with
climatic, tectonic, and eustatic events. Geology 13:637–640
Barry JC, Morgan ME, Winkler AJ, Flynn LJ, Lindsay EH, Jacobs
LL, Pilbeam D (1991) Faunal interchange and Miocene terres-
trial vertebrates of southern Asia. Paleobiology 17:231–245
Berggren WA, Prothero DR (1992) Eocene-Oligocene climatic and
biotic evolution: an overview. In: Berggren WA, Prothero DR
(eds) Eocene–Oligocene climatic and biotic evolution. Princeton
University Press, Princeton, pp 1–28
Bettis EA, Milius AK, Carpenter SJ, Larick R, Zaim Y, Rizal Y,
Ciochon RL, Tassier-Surine SA, Murray D, Bronto S (2009)
Way out of Africa: early Pleistocene paleoenvironments inhab-
ited by Homo erectus in Sangiran, Java. J Hum Evol 56:11–24
Bintanja R, van de Wal SW, Oelemans J (2005) Modelled
atmospheric temperatures and global sea levels of the past
million years. Nature 437:125–128
Bird MI, Taylor D, Hunt C (2005) Palaeoenvironments of insular
Southeast Asia during the last glacial period: a savanna corridor
in Sundaland? Quat Sci Rev 24:2228–2242
Blackburn DC, Bickford DP, Diesmos AC, Iskandar DT, Brown RM
(2010) An ancient origin for the enigmatic flat-headed frogs
(Bombinatoridae: Barbourula) from the islands of Southeast
Asia. PLoS One 5:e12090
Brandon-Jones D (1996) The Asian Colobinae (Mammalia: Cer-
copithecidae) as indicators of quaternary climatic change. Biol J
Linn Soc 59:327–350
Brandon-Jones D (1998) Pre-glacial Bornean primate impoverish-
ment and Wallace’s line. In: Hall R, Holloway JD (eds)
Biogeography and geological evolution of SE Asia. Backhuys,
Leiden, pp 393–403
Brown JW, Van Tuinen M (2011) Evolving perceptions on the
antiquity of the modern avian tree. Living Dinosaurs: the
evolutionary history of modern birds. Wiley, New York,
pp 306–324
Brown JW, Rest JS, Garcıa-Moreno J, Sorenson MD, Mindell D
(2008) Strong mitochondrial DNA support for a Cretaceous
origin of modern avian lineages. BMC Biol 6:6. doi:10.1186/
1741-7007-6-6
Cannon CH, Morley RJ, Bush ABG (2009) The current refugial
rainforests of Sundaland are unrepresentative of their biogeo-
graphic past and highly vulnerable to disturbance. Proc Natl
Acad Sci USA 106:11188–11193
Choi DL-T (1996) Geology of Kinabalu. In: Wong, KM, A Phillipps
(eds) Kinabalu, summit of Borneo. Sabah Society and Sabah
Parks, Kota Kinabalu, Sabah, pp 19–29
Chua VL, Phillipps Q, Lim HC, Taylor SS, Gawin DF, Rahman MA,
Moyle RG, Sheldon FH (2015) Phylogeography of three
endemic birds of Maratua Island, a potential archive of Bornean
biogeography. Raffles Bull Zool (in press)
Cibois A, Kalyakin MV, Han LX, Pasquet E (2002) Molecular
phylogenetics of babblers (Timaliidae): Re-evaluation of the
genera Yuhina and Stachyris. J Avian Biol 33:380–390
Cibois A, Thibault J-C, Bonillo C, Filardi CE, Watling D, Pasquet E
(2014) Phylogeny and biogeography of the fruit doves (Aves:
Columbidae). Mol Phylogenet Evol 70:442–453
Clarke JA, Ksepka DT, Smith NA, Norell MA (2009) Combined
phylogenetic analysis of a new North American fossil species
confirms widespread Eocene distribution for stem rollers (Aves,
Coracii). Zool J Linn Soc 157:586–611
Collar NJ (2004) Species limits in some Indonesian thrushes. Forktail
20:71–87
Cottam M, Hall R, Sperber C, Armstrong R (2010) Pulsed emplace-
ment of the Mount Kinabalu granite, northern Borneo. J Geol
Soc 167:49–60
Cracraft J (2014) Avian higher-level relationships and classification:
Passeriformes. In: Dickinson, EC, L Christidis (eds) The Howard
and Moore complete checklist of the birds of the world, 4th edn,
vol 2. Aves Press, Eastbourne, U.K., pp 17–45
Cranbrook E (2000) Northern Borneo environments of the past 40,000
years. Sarawak Mus J 55:61–109
Cranbrook E (2010) Late Quaternary turnover of mammals in Borneo:
the zooarchaeological record. Biodivers Conserv 19:373–391
Crowe TM, Bowie RCK, Bloomer P, Mandiwana TG, Hedderson
TAJ, Randi E, Wakeling J (2006) Phylogenetics, biogeography
and classification of, and character evolution in, gamebirds
(Aves: Galliformes): effects of character exclusion, data parti-
tioning and missing data. Cladistics 22:1–38
Darlington PJ (1957) Zoogeography: the geographical distribution of
animals. Wiley, New York
Davison GWH (1999) Notes on the taxonomy of some Bornean birds.
Sarawak Mus J 54:289–299
de Bruyn M, Stelbrink B, Morley RJ, Hall R, Carvalho GR, Cannon
CH, van den Bergh G, Meijaard E, Metcalfe I, Boitani L,
Maiorano L, Shoup R, von Rintelen K (2014) Borneo and
Indochina are major evolutionary hotspots for Southeast Asian
biodiversity. Syst Biol 63:879–901
deMenocal PB (2004) African climate change and faunal evolution
during the Pliocene–Pleistocene. Earth Planet Sci Lett 220:3–24
den Tex RJ, Leonard JA (2013) A molecular phylogeny of Asian
barbets: Speciation and extinction in the tropics. Mol Phylogenet
Evol 68:1–13
Dinesen L, Lehmberg T, Svendsen TO, Hansen LA, Fjeldsa J (1994)
A new genus and species of perdicine bird (Phasianidae,
Perdicini) from Tanzania; a relict form with Indo-Malayan
affinities. Ibis 136:3–11
Drummond AJ, Rambaut A (2007) BEAST: Bayesian evolutionary
analysis by sampling trees. BMC Evol Biol 7:214
Drummond AJ, Ho SYW, Phillips MJ, Rambaut A (2006) Relaxed
phylogenetics and dating with confidence. PLoS Biol 4:699–710
Durkee EF (1993) Oil, geology and changing concepts in the
Southwest Philippines (Palawan and the Sulu Sea). Bull Geol
Soc Malays 33:241–262
Ericson PG (2012) Evolution of terrestrial birds in three continents:
biogeography and parallel radiations. J Biogeogr 39:813–824
Ericson PG, Irestedt M, Johansson US (2003) Evolution, biogeogra-
phy, and patterns of diversification in passerine birds. J Avian
Biol 34:3–15
Ericson PGP, Klopstein S, Irestedt M, Nguyen JMT, Nylander JAA
(2014) Dating the divergence of major lineages of Passeriformes.
BMC Evol Biol 14(8):1–15
Esselstyn JA, Oliveros CH, Moyle RG, Peterson AT, McGuire JA,
Brown RM (2010) Integrating phylogenetic and taxonomic
evidence illuminates complex biogeographic patterns along Hux-
ley’s modification of Wallace’s Line. J Biogeogr 37:2054–2066
Fabre PH, Irestedt M, Fjeldsa J, Bristol R, Groombridge JJ, Irham M,
Jønsson KA (2012) Dynamic colonization exchanges between
continents and islands drive diversification in paradise-flycatch-
ers (Terpsiphone, Monarchidae). J Biogeogr 39:1900–1918
J Ornithol
123
Feduccia A (1999) The origin and evolution of birds, 2nd edn. Yale
University Press, New Haven
Feduccia A (2003) ‘Big bang’for tertiary birds? Trends Ecol Evol
18:172–176
Fernando P, Vidya TNC, Payne J, Stuewe M, Davison G, Alfred RJ,
Andau P, Bosi E, Kilbourn A, Melnick DJ (2003) DNA analysis
indicates that Asian elephants are native to Borneo and are
therefore a high priority for conservation. PloS Biol 1:110–115
Fjeldsa J (2013) The global diversification of songbirds (Oscines) and
the build-up of the Sino-Himalayan diversity hotspot. Chin Birds
4:132–143
Fjeldsa J, Bowie RCK (2008) New perspectives on the origin and
diversification of Africa’s forest avifauna. Afr J Ecol
46:235–247
Fuchs J, Fjeldsa J, Bowie RCK, Voelker G, Pasquet E (2006a) The
African warbler genus Hyliota as a lost lineage in the Oscine
songbird tree: molecular support for an African origin of the
Passerida. Mol Phylogenet Evol 39:186–197
Fuchs J, Ohlson JI, Ericson PGP, Pasquet E (2006b) Molecular
phylogeny and biogeographic history of the piculets (Piciformes:
Picumninae). J Avian Biol 37:487–496
Gardner JL, Trueman JW, Ebert D, Joseph L, Magrath RD (2010)
Phylogeny and evolution of the Meliphagoidea, the largest radiation
of Australasian songbirds. Mol Phylogenet Evol 55:1087–1102
Gathorne-Hardy FJ, Syaukani Davies RG, Eggleton P, Jones DT
(2002) Quaternary rainforest refugia in south-east Asia: using
termites (Isoptera) as indicators. Biol J Linn Soc 75:453–466
Gawin DF (2014) Population genetic and hybridization studies of
three Bornean birds species: Mountain Black-eye (Chlorocharis
emiliae), White-rumped Shama (Copsychus malabaricus), and
Oriental Magpie-Robin (Copsychus saularis). In: PhD thesis,
Louisiana State University, Baton Rouge
Gawin DF, Rahman MA, Ramji MFS, Smith BT, Lim HC, Moyle
RG, Sheldon FH (2014) Patterns of avian diversification in
Borneo: the case of the endemic Mountain Black-eye (Chlor-
ocharis emiliae). Auk Adv Ornithol 131:86–99
Gill F, Donsker D (2014) IOC world bird list (v 4.4). Available at:
http://www.worldbirdnames.org/
Gonzalez J-CT (2012) Origin and diversification of hornbills
(Bucerotidae). Oxford University, Oxford
Gonzalez J-CT, Sheldon BC, Collar NJ, Tobias JA (2013) A
comprehensive molecular phylogeny for the hornbills (Aves:
Bucerotidae). Mol Phylogenet Evol 67:468–483
Gorog AJ, Sinaga MH, Engstrom MD (2004) Vicariance or dispersal?
Historical biogeography of three Sunda shelf murine rodents
(Maxomys surifer, Leopoldamys sabanus and Maxomys white-
headi). Biol J Linn Soc 81:91–109
Hall R (1998) The plate tectonics of Cenozoic SE Asia and the
distribution of land and sea. In: Hall R, Holloway JD (eds)
Biogeography and geological evolution of SE Asia. Backhuys,
Leiden, pp 99–131
Hall R (2002) Cenozoic geological and plate tectonic evolution of SE
Asia and the SW Pacific: computer-based reconstructions and
animations. J Asian Earth Sci 20:353–434
Hall R (2009) Southeast Asia’s changing palaeogeography. Blumea
Biodivers Evol Biogeogr Plants 54:1–3
Hall R (2012) Sundaland and Wallacea: geology, plate tectonics and
palaeogeography. In: Gower DJ, Johnson KG, Richardson JE,
Rosen BR, Ruber L, Williams ST (eds) Biotic evolution and
environmental change in Southeast Asia. Cambridge University
Press, Cambridge, pp 32–78
Hall R (2013) The palaeogeography of Sundaland and Wallacea since
the Late Jurassic. J Limnol 72(s2):1–17
Hall R, Nichols G (2002) Cenozoic sedimentation and tectonis in
Borneo: climatic influences on orogenesis. Geol Soc Lond Spec
Publ 191:5–22
Handiani D, Paul A, Prange M, Merkel U, Dupont L, Zhang X (2013)
Tropical vegetation response to Heinrich Event 1 as simulated
with the UVic ESCM and CCSM3. Clim Past 9:1683–1696
Hanebuth T, Stattegger K, Grootes PM (2000) Rapid flooding of the
Sunda Shelf: a late-glacial sea-level record. Science 288:1033–1035
Harzhauser M, Kroh A, Mandic O, Piller WE, Gohlich U, Reuter M,
Berning B (2007) Biogeographic responses to geodynamics: a
key study all around the Oligo-Miocene Tethyan Seaway. Zool
Anz 246:241–256
Heaney LR (1986) Biogeography of mammals in SE Asia: estimates
of rates of colonization, extinction and speciation. Biol J Linn
Soc 28:127–165
Heaney LR (1991) A synopsis of climatic and vegetational change in
Southeast Asia. Clim Change 19:53–61
Hirai H, Wijayanto H, Tanaka H, Mootnick AR, Hayano A,
Perwitasari-Farajallah D, Iskandriati D, Sajuthi D (2005) A
whole-arm translocation (WAT8/9) separating Sumatran and
Bornean agile gibbons, and its evolutionary features. Chromo-
some Res 13:123–133
Ho SY, Phillips MJ (2009) Accounting for calibration uncertainty in
phylogenetic estimation of evolutionary divergence times. Syst
Biol 58:367–380
Holt BG, Lessard J-P, Borregaard MK, Fritz SA, Araujo MB,
Dimitrov D, Fabre P-H, Graham CH, Graves GR, Jønsson KA
(2013) An update of Wallace’s zoogeographic regions of the
world. Science 339:74–78
Hosner PA, Sheldon FH, Lim HC, Moyle RG (2010) Phylogeny and
biogeography of the Asian trogons (Aves: Trogoniformes)
inferred from nuclear and mitochondrial DNA sequences. Mol
Phylogenet Evol 57:1219–1225
Hu J, Pa Peng, Fang D, Jia G, Jian Z, Wang P (2003) No aridity in
Sunda Land during the Last Glaciation: evidence from
molecular-isotopic stratigraphy of long-chain n-alkanes. Palaeo-
geogr Palaeoclimatol Palaeoecol 201:269–281
Hughes JB, Round PD, Woodruff DS (2003) The Indochinese-
Sundaic faunal transition at the Isthmus of Kra: an analysis of
resident forest bird species distributions. J Biogeogr 30:569–580
Irestedt M, Johansson US, Parsons TJ, Ericson PGP (2001) Phylogeny
of major lineages of suboscines (Passeriformes) analysed by
nuclear DNA sequence data. J Avian Biol 32:15–25
Iwanaga H, Teshima KM, Khatab IA, Inomata N, Finkeldey R,
Siregar IZ, Siregar UJ, Szmidt AE (2012) Population structure
and demographic history of a tropical lowland rainforest tree
species Shorea parvifolia (Dipterocarpaceae) from Southeastern
Asia. Ecol Evol 2:1663–1675
James HF (2005) Paleogene fossils and the radiation of modern birds.
Auk 122:1049–1054
Johansson US, Erickson C (2004) A re-evaluation of basal phyloge-
netic relationships within trogons (Aves: Trogonidae) based on
nuclear DNA sequences. J Zool Syst Evol Res 43:166–173
Johansson US, Alstrom P, Olsson U, Ericson PGR, Sundberg P, Price
TD (2007) Build-up of the Himalayan avifauna through immi-
gration: a biogeographical analysis of the Phylloscopus and
Seicercus warblers. Evolution 61:324–333
Jønsson KA, Fjeldsa J (2006) Determining biogeographical patterns
of dispersal and diversification in oscine passerine birds in
Australia, Southeast Asia and Africa. J Biogeogr 33:1155–1165
Jønsson KA, Fjeldsa J, Ericson PGP, Irestedt M (2007) Systematic
placement of an enigmatic Southeast Asian taxon Eupetes
macrocerus and implications for the biogeography of a main
songbird radiation, the Passerida. Biol Lett 3:323–326
Jønsson KA, Irestedt M, Fuchs J, Ericson PGP, Christidis L, Bowie
RCK, Norman JA, Pasquet E, Fjeldsa J (2008) Explosive avian
radiations and multi-directional dispersal across Wallacea:
evidence from the Campephagidae and other Crown Corvida
(Aves). Mol Phylogenet Evol 47:221–236
J Ornithol
123
Jønsson KA, Bowie RCK, Moyle RG, Christidis L, Norman JA, Benz
BW, Fjeldsa J (2010a) Historical biogeography of an Indo-
Pacific passerine bird family (Pachycephalidae): different
colonization patterns in the Indonesian and Melanesian
archipelagos. J Biogeogr 37:245–257
Jønsson KA, Bowie RCK, Moyle RG, Irestedt M, Christidis L,
Norman JA, Fjeldsa J (2010b) Phylogeny and biogeography of
Oriolidae (Aves: Passeriformes). Ecography 33:232–241
Jønsson KA, Irestedt M, Ericson PGP, Fjeldsa J (2010c) A molecular
phylogeny of minivets (Passeriformes: Campephagidae: Peri-
crocotus): implications for biogeography and convergent plu-
mage evolution. Zool Scr 39:1–8
Jønsson KA, Fabre PH, Ricklefs RE, Fjeldsa J (2011) Major global
radiation of corvoid birds originated in the proto-Papuan
archipelago. Proc Natl Acad Sci USA 108:2328–2333
Jønsson KA, Irestedt M, Christidis L, Clegg SM, Holt BG, Fjeldsa J
(2014) Evidence of taxon cycles in an Indo-Pacific passerine bird
radiation (Aves: Pachycephala). Proc R Soc B Biol Sci
281:20131727
Kashiwaya K, Ochiai S, Sakai H, Kawai T (2001) Orbit-related long-
term climate cycles revealed in a 12-Myr continental record from
Lake Baikal. Nature 410:71–74
Kershaw A, Van Der Kaars S, Flenley J (2007) The Quaternary
history of far eastern rainforests. Tropical rainforest responses to
climatic change. Springer-Praxis, Chichester, pp 77–115
Kimball RT, Braun EL, Ligon JD, Lucchini V, Randi E (2001) A
molecular phylogeny of the peacock-pheasants (Galliformes:
Polyplectron spp.) indicates loss and reduction of ornamental
traits and display behaviours. Biol J Linn Soc 73:187–198
Klicka J, Voelker G, Spellman GM (2005) A molecular phylogenetic
analysis of the ‘‘true thrushes’’. Mol Phylogenet Evol
34:486–500
Koufos GD, Kostopoulos DS, Vlachou TD (2005) Neogene/Quater-
nary mammalian migrations in Eastern Mediterranean. Belg J
Zool 135:181–190
Ksepka DT, Clarke JA (2009) Affinities of Palaeospiza bella and the
phylogeny and biogeography of mousbirds (Coliiformes). Auk
126:245–259
Kumar P, Yuan X, Kumar MR, Kind R, Li X, Chadha R (2007) The
rapid drift of the Indian tectonic plate. Nature 449:894–897
Kundu S, Jones CG, Prys-Jones RP, Groombridge JJ (2012) The
evolution of the Indian Ocean parrots (Psittaciformes): extinc-
tion, adaptive radiation and eustacy. Mol Phylogenet Evol
62:296–305
Li Z, Powell CM (2001) An outline of the palaeogeographic evolution
of the Australasian region since the beginning of the Neopro-
terozoic. Earth Sci Rev 53:237–277
Li XW, Walker D (1986) The plant geography of Yunnan Province,
southwest China. J Biogeogr 13:367–397
Li J-T, Li Y, Klaus S, Rao D-Q, Hillis DM, Zhang Y-P (2013)
Diversification of rhacophorid frogs provides evidence for
accelerated faunal exchange between India and Eurasia during
the Oligocene. Proc Natl Acad Sci USA 110:3441–3446
Lim HC, Sheldon FH (2011) Multilocus analysis of the evolutionary
dynamics of rainforest bird populations in Southeast Asia. Mol
Ecol 20:3414–3438
Lim HC, Zou F, Taylor SS, Marks BD, Moyle RG, Voelker G,
Sheldon FH (2010) Phylogeny of magpie-Robins and shamas
(Aves: Turdidae: Copsychus and Trichixos): implications for
island biogeography in Southeast Asia. J Biogeogr
37:1894–1906
Lim HC, Rahman MA, Lim SLH, Moyle RG, Sheldon FH (2011)
Revisiting Wallace’s haunt: coalescent simulations and com-
parative niche modeling reveal historical mechanisms that
promoted avian population divergence in the Malay Archipela-
go. Evolution 65:321–334
Lim HC, Chua VL, Benham PM, Oliveros CH, Rahman MA, Moyle
RG, Sheldon FH (2014) Divergence history of the Rufous-tailed
Tailorbird (Orthotomus sericeus) of Sundaland: implications for
the biogeography of Palawan and the taxonomy of island species
in general. Auk 131:629–642
Lohman DJ, de Bruyn M, Page T, von Rintelen K, Hal R, Ng PKL,
Shih HT, Carvalho GR, von Rintelen T (2011) Biogeography of
the Indo-Australian Archipelago. Annu Rev Ecol Evol Syst
42:205–226
Louys J, Meijaard E (2010) Palaeoecology of Southeast Asian
megafauna-bearing sites from the Pleistocene and a review of
environmental changes in the region. J Biogeogr 37:1432–1449
MacKinnon JR, Phillipps K (1999) A field guide to the birds of Borneo,
Sumatra, Java and Bali. Oxford University Press, Oxford
Mann CF (2008) The birds of Borneo, an annotated checklist. British
Ornithologists’ Union and British Ornithologists’ Club,
Peterborough
Mayr E (1944) Wallace’s line in light of recent zoogeographic
studies. Q Rev Biol 19:1–14
Mayr G (2005) The Paleogene fossil record of birds in Europe. Biol
Rev 80:515–542
Mayr G (2013) The age of the crown group of passerine birds and its
evolutionary significance–molecular calibrations versus the
fossil record. Syst Biodivers 11:7–13
Mayr G (2014) The origins of crown group birds: molecules and
fossils. Palaeontology 57:231–242
Medway L (1972) The Quaternary mammals of Malesia: a review. In:
Ashton P, Ashton M (eds) Transactions of the second Aberdeen-
Hull symposium on Malesian ecology. University of Hull, Hull,
pp 63–83
Medway L, Wells DR (1976) The birds of the Malay Peninsula, vol 5.
H.F. & G. Witherby, London
Mees GF (1986) A list of birds recorded from Bangka Island,
Indonesia. Zool Verh Leiden 232:1–176
Mees GF (1996) Geographical variation in birds of Java. Publ Nuttall
Ornithol Club 26:1–119
Meijaard E (2003) Mammals of south-east Asian islands and their
Late Pleistocene environments. J Biogeogr 30:1245–1257
Meijaard E, Groves C (2004) The biogeographical evolution and
phylogeny of the genus Presbytis. Primate Rep 68:71–90
Meijaard E, van der Zon APM (2003) Mammals of south-east Asian
islands and their Late Pleistocene environments. J Biogeogr
30:1245–1257
Meijer HJM (2014) The avian fossil record in Insular Southeast Asia
and its implications for avian biogeography and palaeoecology.
PeerJ 2:e295
Meulenkamp JE, Sissingh W (2003) Tertiary palaeogeography and
tectonostratigraphic evolution of the Northern and Southern Peri-
Tethys platforms and the intermediate domains of the African-
Eurasian convergent plate boundary zone. Palaeogeogr Palaeo-
climatol Palaeoecol 196:209–228
Mittelbach GG, Schemske DW, Cornell HV, Allen AP, Brown JM,
Bush MB, Harrison SP, Hurlbert AH, Knowlton N, Lessios HA,
McCain CM, McCune AR, McDade LA, McPeek MA, Near TJ,
Price TD, Ricklefs RE, Roy K, Sax DF, Schluter D, Sobel JM,
Turelli M (2007) Evolution and the latitudinal diversity gradient:
speciation, extinction and biogeography. Ecol Lett 10:315–331
Molengraaff GAF (1921) Modern deep sea research in the East Indian
archipelago. Geogr J 57:95–121
Moltesen M, Irestedt M, Fjeldsa J, Ericson PG, Jønsson KA (2012)
Molecular phylogeny of Chloropseidae and Irenidae–Cryptic
species and biogeography. Mol Phylogenet Evol 65:903–914
Morley RJ (1998a) Palynological evidence for Tertiary plant disper-
sals in the SE Asian region in relation to plate tectonic and
climate. In: Hall R, Holloway JD (eds) Biogeography and
geological evolution of SE Asia. Backhuys, Leiden, pp 211–234
J Ornithol
123
Morley RJ (1998b) Tertiary history of the Malesian flora: a
palynological perspective. In: Taxonomy: the cornerstone of
biodiversity: Proc 4th Int Flora Malesiana Symp. Kuala Lumpur,
pp 197–210
Morley RJ (2000) Origin and evolution of tropical rain forests. Wiley,
New York
Morley RJ (2003) Interplate dispersal paths for megathermal
angiosperms. Perspect Plant Ecol Evol Syst 6:5–20
Morley RJ (2011) Cretaceous and Tertiary climate change and the
past distribution of megathermal rainforests. In: Bush M, JR
Flenley (eds) Tropical rainforest responses to climatic change.
Springer SBM, Berlin Heidelberg New York, pp 1–34
Morley RJ (2012) A review of the Cenozoic palaeoclimate history of
Southeast Asia. In: Gower DJ, Johnson KG, Richardson JE,
Rosen BR, Ruber L, Williams ST (eds) Biotic evolution and
environmental change in Southeast Asia. Cambridge University
Press, Cambridge, pp 79–114
Morley RJ, Morley HP (2011) Neogene climate history of the
Makassar Straits. In: The Southeast Asian Gateway: history and
tectonics of Australia-Asia collision. Geological Society of
London, London, pp 319–32
Morley RJ, Morley HP (2013) Mid Cenozoic freshwater wetlands of
the Sunda region. J Limnol 72(s2):18–35
Moss SJ, Wilson MEJ (1998) Biogeographic implications of the
Tertiary palaeogeographic evolution of Sulawesi and Borneo. In:
Hall R, Holloway JD (eds) Biogeography and geological
evolution of SE Asia. Backhuys, Leiden, pp 133–155
Moyle RG (2002) Molecular systematics of barbets and trogons:
pantropical biogeography, African speciation, and issues in
phylogenetic inference. In: PhD thesis, Louisiana State Univer-
sity, Baton Rouge
Moyle RG (2004) Phylogenetics of barbets (Aves: Piciformes) based
on nuclear and mitochondrial DNA sequence data. Mol Phylo-
genet Evol 30:187–200
Moyle RG (2005) Phylogeny and biogeographical history of Trogo-
niformes, a pantropical bird order. Biol J Linn Soc 84:725–738
Moyle RG, Schilthuizen M, Rahman MA, Sheldon FH (2005)
Molecular phylogenetic analysis of the white-crowned forktail
Enicurus leschenaulti in Borneo. J Avian Biol 36:96–101
Moyle RG, Chesser RT, Prum RO, Schikler P, Cracraft J (2006a)
Phylogeny and evolutionary history of Old World suboscine
birds (Aves : Eurylaimides). Am Mus Novit 3544:1–22
Moyle RG, Cracraft J, Lakim M, Nais J, Sheldon FH (2006b)
Reconsideration of the phylogenetic relationships of the enig-
matic Bornean Bristlehead (Pityriasis gymnocephala). Mol
Phylogenet Evol 39:893–898
Moyle RG, Hosner PA, Nais J, Lakim M, Sheldon FH (2008)
Taxonomic status of the Kinabalu ‘linchi’ swiftlet. Bull Br
Ornithol Club 128:94–100
Moyle RG, Filardi CE, Smith CE, Diamond J (2009) Explosive
Pleistocene diversification and hemispheric expansion of a
‘‘great speciator’’. Proc Natl Acad Sci USA 106:1863–1868
Moyle RG, Taylor SS, Oliveros CH, Lim HC, Haines CL, Rahman
MA, Sheldon FH (2011) Diversification of an insular Southeast
Asian genus: Phylogenetic relationships of the spiderhunters
(Aves: Nectariniidae). Auk 128:777–788
Moyle RG, Andersen MJ, Oliveros CH, Steinheimer FD, Reddy S
(2012) Phylogeny and Biogeography of the Core Babblers
(Aves: Timaliidae). Syst Biol 61:631–651
Nesbitt SJ, Ksepka DT, Clarke JA (2011) Podargiform affinities of the
enigmatic Fluvioviridavis platyrhamphus and the early diversi-
fication of Strisores (‘‘Caprimulgiformes’’ plus Apodiformes).
PloS One 6(11)
Newsome J, Flenley J (1988) Late Quaternary vegetational history of
the Central Highlands of Sumatra. II. Palaeopalynology and
vegetational history. J Biogeogr 15:555–578
Ohtani M, Kondo T, Tani N, Ueno S, Lee LS, Ng KKS, Muhammad
N, Finkeldey R, Na’iem M, Indrioko S, Kamiya K, Harada K,
Diway B, Khoo E, Kawamura K, Tsumura Y (2013) Nuclear and
chloroplast DNA phylogeography reveals Pleistocene diver-
gence and subsequent secondary contact of two genetic lineages
of the tropical rainforest tree species Shorea leprosula (Dipte-
rocarpaceae) in South-East Asia. Mol Ecol 22:2264–2279
Oliveros CH, Moyle RG (2010) Origin and diversification of
Philippine bulbuls. Mol Phylogenet Evol 54:822–832
Olson SL (1973) A classification of the Rallidae. Wilson Bull
85:381–416
Olson SL (1979) Picathartes—another West African forest relict with
probable Asian affinities. Bull Br Ornithol Club 99:112–113
Pacheco MA, Battistuzzi FU, Lentino M, Aguilar RF, Kumar S,
Escalante AA (2011) Evolution of modern birds revealed by
mitogenomics: timing the radiation and origin of major orders.
Mol Biol Evol 28:1927–1942
Packert M, Martens J, Sun YH, Severinghaus LL, Nazarenko AA,
Ting J, Topfer T, Tietze DT (2012) Horizontal and elevational
phylogeographic patterns of Himalayan and Southeast Asian
forest passerines (Aves: Passeriformes). J Biogeogr 39:556–573
Patou ML, Wilting A, Gaubert P, Esselstyn JA, Cruaud C, Jennings
AP, Fickel J, Veron G (2010) Evolutionary history of the
Paradoxurus palm civets–a new model for Asian biogeography.
J Biogeogr 37:2077–2097
Phillipps Q, Phillipps K (2014) Phillipps’ field guide to the birds of
Borneo, 3rd edn. John Beaufoy, Oxford
Pook CE, Joger U, Stumpel N, Wuster W (2009) When continents
collide: Phylogeny, historical biogeography and systematics of
the medically important viper genus Echis (Squamata: Serpen-
tes: Viperidae). Mol Phylogenet Evol 53:792–807
Prentice I, Harrison S, Bartlein P (2011) Global vegetation and
terrestrial carbon cycle changes after the last ice age. New Phytol
189:988–998
Quek SP, Davies SJ, Ashton PS, Itino T, Pierce NE (2007) The
geography of diversification in mutualistic ants: a gene’s-eye
view into the Neogene history of Sundaland rain forests. Mol
Ecol 16:2045–2062
Ramji MFS, Rahman MA, Tuen AA (2012) Morphological variation
of Mountain Blackeye (Chlorocharis emiliae) populations in
Malaysian Borneo. Malays Appl Biol 41:1–10
Reddy S (2008) Systematics and biogeography of the shrike-babblers
(Pteruthius): species limits, molecular phylogenetics, and diver-
sification patterns across southern Asia. Mol Phylogenet Evol
47:54–72
Reddy S, Moyle RG (2011) Systematics of the scimitar babblers
(Pomatorhinus: Timaliidae): phylogeny, biogeography, and
species-limits of four species complexes. Biol J Linn Soc
102:846–869
Richardson JE, Costion CM, Muellner AN (2012) The Malesian
floristic interchange: plant migration patterns across Wallace’s
Line. In: Gower DJ, Richardson JE, Rosen BR, Ruber L, Williams
ST (eds) Biotic evolution and environmental change in Southeast
Asia. Cambridge University Press, Cambridge, pp 138–63
Ricklefs RE, Cox GW (1978) Stage of taxon cycle, habitat
distribution, and population density in the avifauna of the West
Indies. Am Nat 112:875–895
Rogl F (1998) Paleogeographic Considerations for Mediterranean and
Paratethys seaways (Oligocene and Miocene). Ann Naturhist
Mus Wien 99A:279–331
Rogl F (1999) Mediterranean and Paratethys. Facts and hypotheses of
an Oligocene to Miocene paleogeography (short overview). Geol
Carpath 50(4):339–349
Round PD, Hughes JB, Woodruff DS (2003) Latitudinal range limits
of resident forest birds in Thailand and the Indochinese-Sundaic
zoogeographic transition. Nat Hist Bull Siam Soc 51:69–96
J Ornithol
123
Ruedi M (1996) Phylogenetic evolution and biogeography of
Southeast Asian shrews (genus Crocidura: Soricidae). Biol J
Linn Soc 58:197–219
Ryan JRJ, Esa YB (2006) Phylogenetic analysis of Hampala fishes
(Subfamily Cyprininae) in Malaysia inferred from partial
mitochondrial Cytochrome b DNA sequences. Zool Sci
23:893–901
Samonds KE, Godfrey LR, Ali JR, Goodman SM, Vences M,
Sutherland MR, Irwin MT, Krause DW (2012) Spatial and
temporal arrival patterns of Madagascar’s vertebrate fauna
explained by distance, ocean currents, and ancestor type. Proc
Natl Acad Sci USA 109:5352–5357
Schweizer M, Seehausen O, Guntert M, Hertwig ST (2010) The
evolutionary diversification of parrots supports a taxon pulse
model with multiple trans-oceanic dispersal events and local
radiations. Mol Phylogenet Evol 54:984–994
Sheldon FH, Moyle RG, Kennard J (2001) Ornithology of Sabah:
history, gazetteer, annotated checklist, and bibliography. Or-
nithol Monogr 52:1–285
Sheldon FH, Lim HC, Nais J, Lakim M, Tuuga A, Malim P, Majuakim J,
Lo A, Schilthuizen M, Hosner PA, Moyle RG (2009a) Observa-
tions on the ecology, distribution, and biogeography of forest birds
in Sabah, Malaysia. Raffles Bull Zool 57:577–586
Sheldon FH, Lohman DJ, Lim HC, Zou F, Goodman SM, Prawiradi-
laga DM, Winker K, Braile TM, Moyle RG (2009b) Phylogeog-
raphy of the magpie-robin species complex (Aves: Turdidae:
Copsychus) reveals a Philippine species, an interesting isolating
barrier, and unusual dispersal patterns in the Indian Ocean and
Southeast Asia. J Biogeogr 36:1070–1083
Sheldon FH, Oliveros CH, Taylor SS, McKay B, Lim HC, Rahman
MA, Mays H, Moyle RG (2012) Molecular phylogeny and
insular biogeography of the lowland tailorbirds of Southeast Asia
(Cisticolidae: Orthotomus). Mol Phylogenet Evol 65:54–63
Siler CD, Oaks JR, Welton LJ, Linkem CW, Swab JC, Diesmos AC,
Brown RM (2012) Did geckos ride the Palawan raft to the
Philippines? J Biogeogr 39:1217–1234
Slik JWF, Raes N, Aiba SI, Brearley FQ, Cannon CH, Meijaard E,
Nagamasu H, Nilus R, Paoli G, Poulsen AD, Sheil D, Suzuki E,
van Valkenburg J, Webb CO, Wilkie P, Wulffraat S (2009)
Environmental correlates for tropical tree diversity and distribu-
tion patterns in Borneo. Divers Distrib 15:523–532
Smythies BE (1999) The Birds of Borneo, 4th edn. Natural History
Publications (Borneo), Kota Kinabalu
Sorenson MD, Payne RB (2005) A molecular genetic analysis of
cuckoo phylogeny. In: Payne RB (ed) The Cuckoos. Oxford
University Press, Oxford, pp 68–94
Stein RW (2013) Multistage scenerios for the evolution of polymor-
phisms in birds. PhD thesis. Department of Biological Sciences.
Simon Fraser University, Burnaby
Steiper ME (2006) Population history, biogeography, and taxonomy
of orangutans (Genus: Pongo) based on a population genetic
meta-analysis of multiple loci. J Hum Evol 50:509–522
Stelbrink B, Albrecht C, Hall R, von Rintelen T (2012) The
biogeography of Sulawesi revisited: is there evidence for a
vicariant origin of taxa on Wallace’s ‘‘anomalous island’’?
Evolution 66:2252–2271
Stuijts IM (1993) Late Pleistocene and Holocene vegetation of West
Java, Indonesia. Mod Quat Res Southeast Asia Southeast Asia
12:1–173
Stuijts I, Newsome J, Flenley J (1988) Evidence for late Quaternary
vegetational change in the Sumatran and Javan highlands. Rev
Palaeobot Palynol 55:207–216
Sun K, Meiklejohn KA, Faircloth BC, Glenn TC, Braun EL, Kimball
RT (2014) The evolution of peafowl and other taxa with ocelli
(eyespots): a phylogenomic approach. Proc R Soc B Biol Sci
281:20140823
van den Bergh GD, de Vos J, Sondaar PY (2001) The Late Quaternary
palaeogeography of mammal evolution in the Indonesian
Archipelago. Palaeogeogr Palaeoclimatol Palaeoecol
171:385–408
van Steenis C (1950) The delimitation of Malaysia and its main plant
geographical divisions. Flora Malesiana 4:70–75
van Tuinen M (2009) Birds (Aves). In: Hedges SB, Kumar S (eds)
The timetree of life. Oxford University Press, New York,
pp 409–411
Viseshakul N, Charoennitikul W, Kitamura S, Kemp A, Thong-Aree
S, Surapunpitak Y, Poonswad P, Ponglikitmongkol M (2011) A
phylogeny of frugivorous hornbills linked to the evolution of
Indian plants within Asian rainforests. J Evol Biol 24:1533–1545
Voelker G, Penalba JV, Huntley JW, Bowie RC (2014) Diversifica-
tion in an Afro-Asian songbird clade (Erythropygia-Copsychus)
reveals founder-event speciation via trans-oceanic dispersals and
a southern to northern colonization pattern in Africa. Mol
Phylogenet Evol 73:97–105
Voris HK (2000) Maps of Pleistocene sea levels in Southeast Asia:
shorelines, river systems and time durations. J Biogeogr
27:1153–1167
Wallace AR (1876) The geographical distribution of animals.
McMillan, London
Wallace AR (1883) The Malay Archipelago. Macmillan, London
Wang N, Kimball RT, Braun EL, Liang B, Zhang Z (2013) Assessing
phylogenetic relationships among Galliformes: a multigene
phylogeny with expanded taxon sampling in Phasianidae. PLoS
One 8(5):e64312
Wells DR (1999) The birds of the Thai-Malay Peninsula, vol 1: Non-
passeries. Academic Press, New York
Wells DR, Dickinson EC, Dekker RWRJ (2003) Systematic notes on
Asian birds. 37. A preliminary review of the Chloropseidae and
Irenidae. Zool Verh Leiden 344:25–42
Whitmore TC (1981) Wallace’s line and plate tectonics. Oxford
University Press, Oxford
Whitmore TC (1987) Biogeographic evolution of the Malay
archipelago. Clarendon Press, Oxford
Whitten T, Soeriaatmadja RE, Afiff SA (1996) The ecology of
Indonesia series, vol II. The Ecology of Java and Bali. Periplus,
Hong Kong
Wilson RCL, Drury SA, Chapman DL (2000) The great Ice age:
climate change and life. Routledge, London
Wilting A, Sollmann R, Meijaard E, Helgen KM, Fickel J (2012)
Mentawai’s endemic, relictual fauna: is it evidence for Pleis-
tocene extinctions on Sumatra? J Biogeogr 39:1608–1620
Witts D, Hall R, Nichols G, Morley R (2012) A new depositional and
provenance model for the Tanjung Formation, Barito Basin, SE
Kalimantan, Indonesia. J Asian Earth Sci 56:77–104
Woodruff DS, Turner LM (2009) The Indochinese-Sundaic zoogeo-
graphic transition: a description and analysis of terrestrial
mammal species distributions. J Biogeogr 36:803–821
Wright TF, Schirtzinger EE, Matsumoto T, Eberhard JR, Graves GR,
Sanchez JJ, Capelli S, Muller H, Scharpegge J, Chambers GK
(2008) A multilocus molecular phylogeny of the parrots
(Psittaciformes): support for a Gondwanan origin during the
Cretaceous. Mol Biol Evol 25:2141–2156
Wurster CM, Bird MI (2014) Barriers and bridges: early human
dispersals in equatorial SE Asia. In: Harff J, Bailey G, Luth F
(eds) Geology and archaeology: submerged landscapes of the
continental shelf. Geological Society, London
Wurster CM, Bird MI, Bull ID, Creed F, Bryant C, Dungait JAJ, Paz
V (2010) Forest contraction in north equatorial Southeast Asia
during the Last Glacial Period. Proc Natl Acad Sci USA
107:15508–15511
Yapp CJ (2004) Fe(CO3)OH in goethite from a mid-latitude North
American Oxisol: estimate of atmospheric CO2 concentration in
J Ornithol
123
the Early Eocene ‘‘climatic optimum’’. Geochim Cosmochim
Acta 68:935–947
Yumul GP, Dimalanta CB, Marquez EJ, Queano KL (2009) Onland
signatures of the Palawan microcontinental block and Philippine
mobile belt collision and crustal growth process: a review.
J Asian Earth Sci 34:610–623
Zachos J, Pagani M, Sloan L, Thomas E, Billups K (2001) Trends,
rhythms, and aberrations in global climate 65 Ma to present.
Science 292:686–693
Zamoros LR, Matsuoka A (2004) Accretion and postaccretion
tectonics of the Calamian Islands, North Palawan block,
Philippines. Island Arc 13:506–619
Zamoros LR, Montes MGA, Queano KL, Marquez EJ, Dimalanta CB,
Gabo JAS, Yumul GP (2008) Buruanga peninsula and Antique
Range: two contrasting terranes in Northwest Panay, Philippines
featuring an arc-continent collision zone. Island Arc 17:443–457
Zhi L, Karesh W, Janczewski D, Frazier-Taylor H, Sajuthi D,
Gombek F, Andau M, Martenson J, O’Brien S (1996) Genomic
differentiation among natural populations of orang-utan (Pongo
pygmaeus). Curr Biol 6:1326–1336
Zhou L, Su YC, Thomas DC, Saunders RM (2012) ‘Out-of-
Africa’dispersal of tropical floras during the Miocene climatic
optimum: evidence from Uvaria (Annonaceae). J Biogeogr
39:322–335
J Ornithol
123
Online Resource 1: List of resident land birds of Sundaland
Article: Return to the Malay Archipelago: the biogeography of Sundaic rainforest birds
Journal: Journal of Ornithology
Authors: Frederick H. Sheldon · Haw Chuan Lim · and Robert G. Moyle
Correspondence: F. H. Sheldon: Museum of Natural Science and Department of Biological Sciences, Louisiana State University,
Baton Rouge, Louisiana, USA; email: [email protected]
Online Resource 1: List of resident land birds of Sundaland, derived from http://avibase.bsc-eoc.org, based on the classification of
Gill and Donsker (2014). Species’ occurrence was also checked using MacKinnon and Phillipps (1999), Phillipps and Phillipps
(2014), Van Marle and Voous (1988), and Wells (1999, 2007). Area abbreviations: B = Borneo, M = Malay Peninsula, S = Sumatra, J
= Java, and P = Palawan; * indicates an island or Malay Peninsula endemic; ** indicates a Sunda endemic
Common Name Scientific Name Area Common Name Scientific Name Area
Megapodes Megapodidae
Drongos Dicruridae
Philippine Megapode Megapodius cumingii B P
Black Drongo Dicrurus macrocercus J
Pheasants & Allies Phasianidae
Ashy Drongo Dicrurus leucophaeus B J M P S
Long-billed Partridge Rhizothera longirostris B M S **
Bronzed Drongo Dicrurus aeneus B M S
Hose's Partridge Rhizothera dulitensis B *
Lesser Racket-tailed Drongo Dicrurus remifer J M S
Black Partridge Melanoperdix niger B M S **
Hair-crested Drongo Dicrurus hottentottus B J P
King Quail Excalfactoria chinensis B J M P S
Sumatran Drongo Dicrurus sumatranus S *
Malaysian Partridge Arborophila campbelli M *
Greater Racket-tailed Drongo Dicrurus paradiseus B J M S
Roll's Partridge Arborophila rolli S *
Fantails Rhipiduridae
Sumatran Partridge Arborophila sumatrana S *
White-throated Fantail Rhipidura albicollis B M S
Grey-breasted Partridge Arborophila orientalis J *
White-bellied Fantail Rhipidura euryura J *
Chestnut-bellied Partridge Arborophila javanica J *
Malaysian Pied Fantail Rhipidura javanica B J M S
Red-billed Partridge Arborophila rubrirostris S *
Philippine Pied Fantail Rhipidura nigritorquis P
Red-breasted Hill Partridge Arborophila hyperythra B *
Spotted Fantail Rhipidura perlata B M S **
Chestnut-necklaced Partridge Arborophila charltonii B M S **
Rufous-tailed Fantail Rhipidura phoenicura J *
Ferruginous Partridge Caloperdix oculeus B M S
Monarchs Monarchidae
Crimson-headed Partridge Haematortyx sanguiniceps B *
Black-naped Monarch Hypothymis azurea B J M P S
Crested Partridge Rollulus rouloul B M S
Asian Paradise Flycatcher Terpsiphone paradisi B J M S
Red Junglefowl Gallus gallus J M S
Japanese Paradise Flycatcher Terpsiphone atrocaudata P
Green Junglefowl Gallus varius J
Blue Paradise Flycatcher Terpsiphone cyanescens P *
Hoogerwerf's Pheasant Lophura hoogerwerfi S *
Crows & Jays Corvidae
Salvadori's Pheasant Lophura inornata S *
Crested Jay Platylophus galericulatus B J M S **
Crestless Fireback Lophura erythrophthalma B M S **
Black Magpie Platysmurus leucopterus B M S **
Crested Fireback Lophura ignita B M S **
Common Green Magpie Cissa chinensis B M S
Bulwer's Pheasant Lophura bulweri B *
Javan Green Magpie Cissa thalassina J *
Bronze-tailed Peacock-Pheasant Polyplectron chalcurum S *
Bornean Green Magpie Cissa jefferyi B *
Mountain Peacock-Pheasant Polyplectron inopinatum M *
Sumatran Treepie Dendrocitta occipitalis S *
Malayan Peacock-Pheasant Polyplectron malacense M *
Bornean Treepie Dendrocitta cinerascens B *
Bornean Peacock-Pheasant Polyplectron schleiermacheri B *
Racket-tailed Treepie Crypsirina temia J M S
Palawan Peacock-Pheasant Polyplectron napoleonis P *
Slender-billed Crow Corvus enca B J M P S
Crested Argus Rheinardia ocellata M
Large-billed Crow Corvus macrorhynchos J M P S
Great Argus Argusianus argus B M S
Rail-babbler Eupetidae
Green Peafowl Pavo muticus J M
Rail-babbler Eupetes macrocerus B M S **
Ospreys Pandionidae
Fairy Flycatchers Stenostiridae
Western Osprey Pandion haliaetus M
Grey-headed Canary-Flycatcher Culicicapa ceylonensis B J M S
Eastern Osprey Pandion cristatus J
Citrine Canary-flycatcher Culicicapa helianthea P
Kites, Hawks & Eagles Accitripidae
Tits Paridae
Black-winged Kite Elanus caeruleus B J M P
Sultan Tit Melanochlora sultanea M S
Crested Honey Buzzard Pernis ptilorhynchus B J M P
Palawan Tit Pardaliparus amabilis P *
Philippine Honey Buzzard Pernis steerei P
Great Tit Parus major B
Jerdon's Baza Aviceda jerdoni B M P
Cinereous Tit Parus cinereus J M S
White-rumped Vulture Gyps bengalensis M
Larks Alaudidae
Red-headed Vulture Sarcogyps calvus M
Horsfield's Bush Lark Mirafra javanica J
Crested Serpent Eagle Spilornis cheela B J M P S
Bulbuls Pycnonotidae
Kinabalu Serpent Eagle Spilornis kinabaluensis B *
Straw-headed Bulbul Pycnonotus zeylanicus B J M S **
Philippine Serpent Eagle Spilornis holospilus P
Cream-striped Bulbul Pycnonotus leucogrammicus S *
Bat Hawk Macheiramphus alcinus B M S
Spot-necked Bulbul Pycnonotus tympanistrigus S *
Changeable Hawk-Eagle Nisaetus cirrhatus B J M P S
Black-and-white Bulbul Pycnonotus melanoleucos B M S **
Mountain Hawk-Eagle Nisaetus nipalensis M
Black-headed Bulbul Pycnonotus atriceps B J M P S
Blyth's Hawk-Eagle Nisaetus alboniger B M S **
Black-crested Bulbul Pycnonotus flaviventris M
Javan Hawk-Eagle Nisaetus bartelsi J *
Ruby-throated Bulbul Pycnonotus dispar J S **
Pinsker's Hawk-Eagle Nisaetus pinskeri P
Bornean Bulbul Pycnonotus montis B *
Wallace's Hawk-Eagle Nisaetus nanus B M S **
Scaly-breasted Bulbul Pycnonotus squamatus B J M S **
Rufous-bellied Hawk-Eagle Lophotriorchis kienerii B J M P S
Grey-bellied Bulbul Pycnonotus cyaniventris B M S **
Indian Black Eagle Ictinaetus malaiensis B J M S
Red-whiskered Bulbul Pycnonotus jocosus M
Greater Spotted Eagle Clanga clanga M S
Sooty-headed Bulbul Pycnonotus aurigaster J *
Crested Goshawk Accipiter trivirgatus B J M P S
Puff-backed Bulbul Pycnonotus eutilotus B M S **
Besra Accipiter virgatus B J
Blue-wattled Bulbul Pycnonotus nieuwenhuisii B S
Brahminy Kite Haliastur indus B J M P S
Orange-spotted Bulbul Pycnonotus bimaculatus J S
White-bellied Sea Eagle Haliaeetus leucogaster B J M P S
Stripe-throated Bulbul Pycnonotus finlaysoni M
Lesser Fish Eagle Haliaeetus humilis B M S
Flavescent Bulbul Pycnonotus flavescens B
Grey-headed Fish Eagle Haliaeetus ichthyaetus B J M P S
Yellow-vented Bulbul Pycnonotus goiavier B J M P S
Rufous-winged Buzzard Butastur liventer J
Olive-winged Bulbul Pycnonotus plumosus B J M S **
Buttonquail Turnicidae
Ashy-fronted Bulbul Pycnonotus cinereifrons P *
Common Buttonquail Turnix sylvaticus J
Streak-eared Bulbul Pycnonotus blanfordi M
Barred Buttonquail Turnix suscitator J M S
Cream-vented Bulbul Pycnonotus simplex B J M S **
Pigeons & Doves Columbidae
Asian Red-eyed Bulbul Pycnonotus brunneus B J M S **
Silvery Pigeon Columba argentina B S
Spectacled Bulbul Pycnonotus erythropthalmos B M S **
Metallic Pigeon Columba vitiensis B P
Finsch's Bulbul Alophoixus finschii B M S **
Island Collared Dove Streptopelia bitorquata J P
Ochraceous Bulbul Alophoixus ochraceus B M S
Spotted Dove Spilopelia chinensis J M P S
Grey-cheeked Bulbul Alophoixus bres B J M S **
Barred Cuckoo-Dove Macropygia unchall J M S
Palawan Bulbul Alophoixus frater P *
Philippine Cuckoo-Dove Macropygia tenuirostris P
Yellow-bellied Bulbul Alophoixus phaeocephalus B M S **
Ruddy Cuckoo-Dove Macropygia emiliana B J S
Hook-billed Bulbul Setornis criniger B S
Little Cuckoo-Dove Macropygia ruficeps B J M S
Hairy-backed Bulbul Tricholestes criniger B M S **
Common Emerald Dove Chalcophaps indica B J M P S
Buff-vented Bulbul Iole olivacea B M S **
Zebra Dove Geopelia striata B J M S
Sulphur-bellied Bulbul Iole palawanensis P *
Nicobar Pigeon Caloenas nicobarica B J M P S
Mountain Bulbul Ixos mcclellandii M
White-eared Brown Dove Phapitreron leucotis P
Streaked Bulbul Ixos malaccensis B M S **
Cinnamon-headed Green Pigeon Treron fulvicollis B M S **
Sunda Bulbul Ixos virescens J S **
Little Green Pigeon Treron olax B J M S ** Cinereous Bulbul Hemixos cinereus B M S
Pink-necked Green Pigeon Treron vernans B J M P S
Swallows & Martins Hirundinidae
Orange-breasted Green Pigeon Treron bicinctus J M S
Pacific Swallow Hirundo tahitica B J M P S
Philippine Green Pigeon Treron axillaris P
Dusky Crag Martin Ptyonoprogne concolor M
Thick-billed Green Pigeon Treron curvirostra B J M P S
Striated Swallow Cecropis striolata J M P
Grey-cheeked Green Pigeon Treron griseicauda J
Wren-babblers Pnoepygidae
Large Green Pigeon Treron capellei B J M S ** Pygmy Wren-babbler Pnoepyga pusilla J M S
Sumatran Green Pigeon Treron oxyurus J S **
Cettia Bush Warblers Cettiidae
Yellow-vented Green Pigeon Treron seimundi M
Yellow-bellied Warbler Abroscopus superciliaris B J M S
Wedge-tailed Green Pigeon Treron sphenurus J M S
Mountain Tailorbird Phyllergates cuculatus B J M P S
Banded Fruit Dove Ptilinopus cinctus J
Sunda Bush Warbler Horornis vulcanius B J P S
Pink-headed Fruit Dove Ptilinopus porphyreus J S
Javan Tesia Tesia superciliaris J *
Yellow-breasted Fruit Dove Ptilinopus occipitalis P
Bornean Stubtail Urosphena whiteheadi B *
Jambu Fruit Dove Ptilinopus jambu B J M S ** Bushtits Aegithalidae
Black-chinned Fruit Dove Ptilinopus leclancheri P
Pygmy Bushtit Psaltria exilis J *
Black-naped Fruit Dove Ptilinopus melanospilus B P
Leaf Warblers & Allies Phylloscopidae
Green Imperial Pigeon Ducula aenea B J M P S
Philippine Leaf Warbler Phylloscopus olivaceus P
Grey Imperial Pigeon Ducula pickeringii B P
Mountain Leaf Warbler Phylloscopus trivirgatus B J M S
Mountain Imperial Pigeon Ducula badia B J M S
Negros Leaf Warbler Phylloscopus nigrorum P
Dark-backed Imperial Pigeon Ducula lacernulata J
Chestnut-crowned Warbler Seicercus castaniceps M S
Pied Imperial Pigeon Ducula bicolor B J M P S
Yellow-breasted Warbler Seicercus montis B M P S
Cuckoos Cuculidae
Sunda Warbler Seicercus grammiceps J S
Short-toed Coucal Centropus rectunguis B M S **
Reed Warblers Acrocephalidae
Sunda Coucal Centropus nigrorufus J *
Oriental Reed Warbler Acrocephalus orientalis J
Greater Coucal Centropus sinensis B J M P S
Clamorous Reed Warbler Acrocephalus stentoreus B J
Lesser Coucal Centropus bengalensis B J M P S
Grassbirds & Allies Locustellidae
Bornean Ground Cuckoo Carpococcyx radiceus B *
Javan Bush Warbler Locustella montis J *
Sumatran Ground Cuckoo Carpococcyx viridis S *
Friendly Bush Warbler Locustella accentor B *
Raffles's Malkoha Rhinortha chlorophaea B M S
Striated Grassbird Megalurus palustris B P
Red-billed Malkoha Zanclostomus javanicus B J M S
Cisticolas & Allies Cisticolidae
Chestnut-breasted Malkoha Phaenicophaeus curvirostris B J M P S
Zitting Cisticola Cisticola juncidis J M P S
Chestnut-bellied Malkoha Phaenicophaeus sumatranus B M S **
Golden-headed Cisticola Cisticola exilis B J P
Black-bellied Malkoha Phaenicophaeus diardi B M S
Brown Prinia Prinia polychroa J
Green-billed Malkoha Phaenicophaeus tristis J M S
Hill Prinia Prinia superciliaris M
Chestnut-winged Cuckoo Clamator coromandus P
Rufescent Prinia Prinia rufescens M
Asian Koel Eudynamys scolopaceus M P S
Bar-winged Prinia Prinia familiaris J S
Asian Emerald Cuckoo Chrysococcyx maculatus S
Yellow-bellied Prinia Prinia flaviventris B J M S
Violet Cuckoo Chrysococcyx xanthorhynchus B J M P S
Plain Prinia Prinia inornata J
Little Bronze Cuckoo Chrysococcyx minutillus B J M S
Common Tailorbird Orthotomus sutorius J M
Banded Bay Cuckoo Cacomantis sonneratii B J M P S
Dark-necked Tailorbird Orthotomus atrogularis B J M S
Plaintive Cuckoo Cacomantis merulinus B J M P S
Rufous-tailed Tailorbird Orthotomus sericeus B M P S **
Rusty-breasted Cuckoo Cacomantis sepulcralis J M P S
Ashy Tailorbird Orthotomus ruficeps B J M S
Square-tailed Drongo-Cuckoo Surniculus lugubris B J M P S
Olive-backed Tailorbird Orthotomus sepium J
Moustached Hawk-Cuckoo Hierococcyx vagans B M S
Babblers Timaliidae
Dark Hawk-Cuckoo Hierococcyx bocki B M S **
Large Scimitar Babbler Pomatorhinus hypoleucos M
Philippine Hawk-Cuckoo Hierococcyx pectoralis P
Chestnut-backed Scimitar Babbler Pomatorhinus montanus B J M S **
Malaysian Hawk-Cuckoo Hierococcyx fugax J M S
White-breasted Babbler Stachyris grammiceps J *
Indian Cuckoo Cuculus micropterus B J M S
Grey-throated Babbler Stachyris nigriceps B M S
Sunda Cuckoo Cuculus lepidus J M S
Grey-headed Babbler Stachyris poliocephala B M S **
Barn Owls Tytonidae
Spot-necked Babbler Stachyris strialata S
Western Barn Owl Tyto alba J M S
Chestnut-rumped Babbler Stachyris maculata B M S **
Grass Owl Tyto capensis B
White-necked Babbler Stachyris leucotis B M S **
Eastern Grass Owl Tyto longimembris P
Black-throated Babbler Stachyris nigricollis B M S **
Oriental Bay Owl Phodilus badius B J M S
White-bibbed Babbler Stachyris thoracica J *
Owls Strigidae
Chestnut-winged Babbler Stachyris erythroptera B M S **
White-fronted Scops Owl Otus sagittatus M
Crescent-chested Babbler Stachyris melanothorax J *
Reddish Scops Owl Otus rufescens B J M S ** Rufous-fronted Babbler Stachyridopsis rufifrons B M S
Mountain Scops Owl Otus spilocephalus B M S
Golden Babbler Stachyridopsis chrysaea M S
Rajah Scops Owl Otus brookii B S
Pin-striped Tit-Babbler Macronus gularis B M P S
Javan Scops Owl Otus angelinae J *
Bold-striped Tit-Babbler Macronus bornensis J M
Mentawai Scops Owl Otus mentawi S *
Grey-cheeked Tit-Babbler Macronus flavicollis J *
Collared Scops Owl Otus bakkamoena B
Fluffy-backed Tit-Babbler Macronus ptilosus B M S **
Sunda Scops Owl Otus lempiji J M S
Chestnut-capped Babbler Timalia pileata J
Palawan Scops Owl Otus fuliginosus P *
Fulvettas & Ground Babblers Pellorneidae
Oriental Scops Owl Otus sunia B M
Rufous-winged Fulvetta Alcippe castaneceps M
Mantanani Scops Owl Otus mantananensis B P **
Brown Fulvetta Alcippe brunneicauda B M S **
Simeulue Scops Owl Otus umbra S *
Javan Fulvetta Alcippe pyrrhoptera J *
Enggano Scops Owl Otus enganensis S *
Mountain Fulvetta Alcippe peracensis M
Barred Eagle-Owl Bubo sumatranus B J M S
Bornean Wren-Babbler Ptilocichla leucogrammica B *
Brown Fish Owl Ketupa zeylonensis M
Falcated Wren-Babbler Ptilocichla falcata P
Buffy Fish Owl Ketupa ketupu B J M S
Rusty-breasted Wren-Babbler Napothera rufipectus S *
Spotted Wood Owl Strix seloputo J M P S
Black-throated Wren-Babbler Napothera atrigularis B *
Brown Wood Owl Strix leptogrammica B J M S
Large Wren-Babbler Napothera macrodactyla J M S **
Collared Owlet Glaucidium brodiei B M S
Marbled Wren-Babbler Napothera marmorata M S **
Javan Owlet Glaucidium castanopterum J *
Streaked Wren-Babbler Napothera brevicaudata M
Brown Hawk-Owl Ninox scutulata B J M P S
Mountain Wren-Babbler Napothera crassa B *
Frogmouths Podargidae
Eyebrowed Wren-Babbler Napothera epilepidota B J M S
Large Frogmouth Batrachostomus auritus B M S **
Collared Babbler Gampsorhynchus torquatus M
Dulit Frogmouth Batrachostomus harterti B *
Sumatran Wren-Babbler Rimator albostriatus S *
Gould's Frogmouth Batrachostomus stellatus B M S **
Abbott's Babbler Malacocincla abbotti B J M S
Short-tailed Frogmouth Batrachostomus poliolophus S *
Horsfield's Babbler Malacocincla sepiaria B J M S **
Bornean Frogmouth Batrachostomus mixtus B *
Black-browed Babbler Malacocincla perspicillata B *
Javan Frogmouth Batrachostomus javensis J *
Short-tailed Babbler Malacocincla malaccensis B M S **
Blyth's Frogmouth Batrachostomus affinis B J M S
Ashy-headed Babbler Malacocincla cinereiceps P *
Palawan Frogmouth Batrachostomus chaseni B P
Moustached Babbler Malacopteron magnirostre B M S **
Sunda Frogmouth Batrachostomus cornutus B M S
Sooty-capped Babbler Malacopteron affine B M S **
Nightjars Caprimulgidae
Scaly-crowned Babbler Malacopteron cinereum B J M S
Malaysian Eared Nightjar Lyncornis temminckii B M S **
Rufous-crowned Babbler Malacopteron magnum B M S **
Great Eared Nightjar Lyncornis macrotis M S
Melodious Babbler Malacopteron palawanense P *
Large-tailed Nightjar Caprimulgus macrurus B J M P S
Grey-breasted Babbler Malacopteron albogulare B M S **
Philippine Nightjar Caprimulgus manillensis P
White-chested Babbler Trichastoma rostratum B M S **
Savanna Nightjar Caprimulgus affinis B J M S
Ferruginous Babbler Trichastoma bicolor B M S **
Bonaparte's Nightjar Caprimulgus concretus B S
Striped Wren-Babbler Kenopia striata B M S **
Salvadori's Nightjar Caprimulgus pulchellus J S
Puff-throated Babbler Pellorneum ruficeps M
Treeswifts Hemiprocnidae
Buff-breasted Babbler Pellorneum tickelli M
Grey-rumped Treeswift Hemiprocne longipennis B J M S
Sumatran Babbler Pellorneum buettikoferi S *
Whiskered Treeswift Hemiprocne comata B M S
Temminck's Babbler Pellorneum pyrrogenys B J S
Swifts Apodidae
Black-capped Babbler Pellorneum capistratum B J M S **
Giant Swiftlet Hydrochous gigas J M S **
Laughingthrushes Leiothrichidae
Glossy Swiftlet Collocalia esculenta B J M S
Sumatran Laughingthrush Garrulax bicolor S *
Cave Swiftlet Collocalia linchi J S
Sunda Laughingthrush Garrulax palliatus B S **
Bornean Swiftlet Collocalia dodgei B *
Rufous-fronted Laughingthrush Garrulax rufifrons J *
Pygmy Swiftlet Collocalia troglodytes P
Chestnut-capped Laughingthrush Garrulax mitratus M S **
Philippine Swiftlet Aerodramus mearnsi P
Chestnut-hooded Laughingthrush Garrulax treacheri B *
Volcano Swiftlet Aerodramus vulcanorum J *
Black Laughingthrush Garrulax lugubris M S **
Mossy-nest Swiftlet Aerodramus salangana B J S
Bare-headed Laughingthrush Garrulax calvus B *
Uniform Swiftlet Aerodramus vanikorensis B P
Malayan Laughingthrush Trochalopteron peninsulae M
Black-nest Swiftlet Aerodramus maximus B J M S
Himalayan Cutia Cutia nipalensis M
Edible-nest Swiftlet Aerodramus fuciphagus B J M
Blue-winged Minla Minla cyanouroptera M
Germain's Swiftlet Aerodramus germani M P
Bar-throated Minla Minla strigula M
Silver-rumped Spinetail Rhaphidura leucopygialis B J M S ** Silver-eared Mesia Leiothrix argentauris M S
White-throated Needletail Hirundapus caudacutus M
Spotted Crocias Crocias albonotatus J *
Brown-backed Needletail Hirundapus giganteus B J M P S
Long-tailed Sibia Heterophasia picaoides M S
Asian Palm Swift Cypsiurus balasiensis B J M P S
White-eyes Zosteropidae
Pacific Swift Apus pacificus J M P
Chestnut-crested Yuhina Yuhina everetti B *
House Swift Apus nipalensis B J M S
Palawan Striped Babbler Zosterornis hypogrammicus P
Trogons Trogonidae
Mees's White-eye Lophozosterops javanicus J *
Javan Trogon Apalharpactes reinwardtii J *
Pygmy White-eye Oculocincta squamifrons B *
Sumatran Trogon Apalharpactes mackloti S *
Mountain Black-eye Chlorocharis emiliae B *
Red-naped Trogon Harpactes kasumba B M S **
Oriental White-eye Zosterops palpebrosus B J M S
Diard's Trogon Harpactes diardii B M S **
Enggano White-eye Zosterops salvadorii S *
Whitehead's Trogon Harpactes whiteheadi B *
Black-capped White-eye Zosterops atricapilla B S
Cinnamon-rumped Trogon Harpactes orrhophaeus B M S **
Everett's White-eye Zosterops everetti B M
Scarlet-rumped Trogon Harpactes duvaucelii B M S **
Yellowish White-eye Zosterops nigrorum P
Orange-breasted Trogon Harpactes oreskios B J M S
Mountain White-eye Zosterops montanus J P S
Red-headed Trogon Harpactes erythrocephalus M S
Javan White-eye Zosterops flavus B J
Rollers Coraciidae
Lemon-bellied White-eye Zosterops chloris J
Indian Roller Coracias benghalensis M
Fairy-bluebirds Irenidae
Oriental Dollarbird Eurystomus orientalis B J M P S
Asian Fairy-bluebird Irena puella B J M P S
Kingfishers Alcedinidae
Nuthatches Sittidae
Rufous-collared Kingfisher Actenoides concretus B M S **
Velvet-fronted Nuthatch Sitta frontalis B J M P S
Banded Kingfisher Lacedo pulchella B J M S
Blue Nuthatch Sitta azurea J M S * *
Stork-billed Kingfisher Pelargopsis capensis B J M P S
Starling Sturnidae
Brown-winged Kingfisher Pelargopsis amauroptera M
Asian Glossy Starling Aplonis panayensis B J M P S
Ruddy Kingfisher Halcyon coromanda B J M P
Short-tailed Starling Aplonis minor J
White-throated Kingfisher Halcyon smyrnensis M P
Coleto Sarcops calvus P
Javan Kingfisher Halcyon cyanoventris J *
Golden-crested Myna Ampeliceps coronatus M
Collared Kingfisher Todiramphus chloris B J M P S
Common Hill Myna Gracula religiosa B J M P S
Cerulean Kingfisher Alcedo coerulescens J S
Nias Hill Myna Gracula robusta S *
Blue-banded Kingfisher Alcedo euryzona B J M S
Enggano Hill Myna Gracula enganensis S *
Blue-eared Kingfisher Alcedo meninting B J M P S
Javan Myna Acridotheres javanicus J *
Common Kingfisher Alcedo atthis J M
Jungle Myna Acridotheres fuscus M
Oriental Dwarf Kingfisher Ceyx erithaca B J M P S
Common Myna Acridotheres tristis M
Bee-eaters Meropidae
Black-winged Starling Acridotheres melanopterus J *
Red-bearded Bee-eater Nyctyornis amictus B M S
Pied Myna Gracupica contra J S
Blue-tailed Bee-eater Merops philippinus J M P S
Daurian Starling Agropsar sturninus J M
Blue-throated Bee-eater Merops viridis B J M P S
Chestnut-cheeked Starling Agropsar philippensis P
Chestnut-headed Bee-eater Merops leschenaulti J M S
White-shouldered Starling Sturnia sinensis M
Hoopoes Upupidae
Bali Myna Leucopsar rothschildi J *
Eurasian Hoopoe Upupa epops M
Thrushes Turdidae
Hornbills Bucerotidae
Chestnut-capped Thrush Geokichla interpres B J M S
White-crowned Hornbill Berenicornis comatus B M S
Enggano Thrush Geokichla leucolaema S *
Rhinoceros Hornbill Buceros rhinoceros B J M S ** Orange-headed Thrush Geokichla citrina B J M S
Great Hornbill Buceros bicornis M S
Siberian Thrush Geokichla sibirica M
Helmeted Hornbill Rhinoplax vigil B M S **
Everett's Thrush Zoothera everetti B *
Palawan Hornbill Anthracoceros marchei P *
Sunda Thrush Zoothera andromedae J S
Oriental Pied Hornbill Anthracoceros albirostris B J M S
Scaly Thrush Zoothera dauma J S
Black Hornbill Anthracoceros malayanus B M S **
Island Thrush Turdus poliocephalus B J S
Bushy-crested Hornbill Anorrhinus galeritus B M S **
Eyebrowed Thrush Turdus obscurus M
Wreathed Hornbill Rhyticeros undulatus B J M S
Sumatran Cochoa Cochoa beccarii S *
Plain-pouched Hornbill Rhyticeros subruficollis M S
Javan Cochoa Cochoa azurea J *
Wrinkled Hornbill Rhabdotorrhinus corrugatus B M S **
Fruithunter Chlamydochaera jefferyi B *
Asian Barbets Megalaimidae
Oriental Magpie-Robin Copsychus saularis B J M S
Fire-tufted Barbet Psilopogon pyrolophus M S **
Rufous-tailed Shama Copsychus pyrropygus B M S **
Lineated Barbet Megalaima lineata J M
Philippine Magpie-Robin Copsychus mindanensis P
Brown-throated Barbet Megalaima corvina J *
White-rumped Shama Copsychus malabaricus J M S
Golden-whiskered Barbet Megalaima chrysopogon B M S **
White-crowned Shama Copsychus stricklandii B *
Red-crowned Barbet Megalaima rafflesii B M S **
White-vented Shama Copsychus niger P *
Red-throated Barbet Megalaima mystacophanos B M S
Old World Flycatchers Muscicapidae
Black-banded Barbet Megalaima javensis J *
Grey-streaked Flycatcher Muscicapa griseisticta B S
Golden-throated Barbet Megalaima franklinii M
Asian Brown Flycatcher Muscicapa latirostris B J
Black-browed Barbet Megalaima oorti M S
Brown-streaked Flycatcher Muscicapa williamsoni M S
Mountain Barbet Megalaima monticola B *
Rufous-browed Flycatcher Anthipes solitaris M S
Yellow-crowned Barbet Megalaima henricii B M S **
Pale Blue Flycatcher Cyornis unicolor B J M S
Flame-fronted Barbet Megalaima armillaris J *
Rück's Blue Flycatcher Cyornis ruckii S *
Golden-naped Barbet Megalaima pulcherrima B *
Hill Blue Flycatcher Cyornis banyumas B J M S
Blue-eared Barbet Megalaima australis B J M S
Large Blue Flycatcher Cyornis magnirostris M
Bornean Barbet Megalaima eximia B *
Palawan Blue Flycatcher Cyornis lemprieri P *
Coppersmith Barbet Megalaima haemacephala J M P S
Tickell's Blue Flycatcher Cyornis tickelliae M S
Brown Barbet Caloramphus fuliginosus B *
Sunda Blue Flycatcher Cyornis caerulatus B S
Sooty Barbet Caloramphus hayii M S **
Bornean Blue Flycatcher Cyornis superbus B *
Honeyguides Indicatoridae
Chinese Blue Flycatcher Cyornis glaucicomans M
Malaysian Honeyguide Indicator archipelagicus B M S
Malaysian Blue Flycatcher Cyornis turcosus B M S **
Woodpeckers Picidae
Mangrove Blue Flycatcher Cyornis rufigastra B J M P S
Speckled Piculet Picumnus innominatus B M S
White-tailed Flycatcher Cyornis concretus B M S
Rufous Piculet Sasia abnormis B J M S ** Fulvous-chested Jungle Flycatcher Cyornis olivaceus B J S
Grey-and-buff Woodpecker Hemicircus concretus B J M S ** Grey-chested Jungle Flycatcher Cyornis umbratilis B J M S **
Sunda Pygmy Woodpecker Dendrocopos moluccensis B J M S
Rufous-tailed Jungle Flycatcher Cyornis ruficauda B
Grey-capped Pygmy Woodpecker Dendrocopos canicapillus B M S
Rufous-vented Niltava Niltava sumatrana M S **
Freckle-breasted Woodpecker Dendrocopos analis J S
Large Niltava Niltava grandis M S
White-bellied Woodpecker Dryocopus javensis B J M P S
Verditer Flycatcher Eumyias thalassinus B M S
Banded Woodpecker Chrysophlegma miniaceum B J M S
Indigo Flycatcher Eumyias indigo B J S
Checker-throated Woodpecker Chrysophlegma mentale B J M S ** Lesser Shortwing Brachypteryx leucophris J M S
Greater Yellownape Chrysophlegma flavinucha M S
White-browed Shortwing Brachypteryx montana B J M P S
Lesser Yellownape Picus chlorolophus M S
Eyebrowed Jungle Flycatcher Vauriella gularis B *
Crimson-winged Woodpecker Picus puniceus B J M S
Siberian Blue Robin Larvivora cyane M
Streak-breasted Woodpecker Picus viridanus M
White-tailed Robin Myiomela leucura M
Laced Woodpecker Picus vittatus J M S
Sunda Robin Myiomela diana J S
Grey-headed Woodpecker Picus canus M S
Sunda Forktail Enicurus velatus J S
Olive-backed Woodpecker Dinopium rafflesii B M S
Chestnut-naped Forktail Enicurus ruficapillus B M S **
Common Flameback Dinopium javanense B J M S
Slaty-backed Forktail Enicurus schistaceus M
Spot-throated Flameback Dinopium everetti P
White-crowned Forktail Enicurus leschenaulti B J M S
Greater Flame-backed Woodpecker Chrysocolaptes lucidus B
Bornean Forktail Enicurus borneensis B *
Red-headed Flameback Chrysocolaptes
erythrocephalus P
Shiny Whistling Thrush Myophonus melanurus S *
Javan Flameback Chrysocolaptes strictus J *
Javan Whistling Thrush Myophonus glaucinus J *
Greater Flameback Chrysocolaptes guttacristatus J M S
Bornean Whistling Thrush Myophonus borneensis B *
Bamboo Woodpecker Gecinulus viridis M
Brown-winged Whistling Thrush Myophonus castaneus S *
Maroon Woodpecker Blythipicus rubiginosus B M S **
Malayan Whistling Thrush Myophonus robinsoni M *
Orange-backed Woodpecker Reinwardtipicus validus B J M S ** Blue Whistling Thrush Myophonus caeruleus J M S
Rufous Woodpecker Micropternus brachyurus B J M S
Rufous-chested Flycatcher Ficedula dumetoria B J M S
Buff-rumped Woodpecker Meiglyptes tristis B J M S
Taiga Flycatcher Ficedula albicilla M
Buff-necked Woodpecker Meiglyptes tukki B M S **
Snowy-browed Flycatcher Ficedula hyperythra B J M P S
Great Slaty Woodpecker Mulleripicus pulverulentus B J M P S
Palawan Flycatcher Ficedula platenae P *
Falcons Falconidae
Little Pied Flycatcher Ficedula westermanni B J M P S
Black-thighed Falconet Microhierax fringillarius B J M S
Pygmy Flycatcher Muscicapella hodgsoni B M S
White-fronted Falconet Microhierax latifrons B *
Blue Rock Thrush Monticola solitarius M P S
Spotted Kestrel Falco moluccensis J
Pied Bush Chat Saxicola caprata J
Peregrine Falcon Falco peregrinus B J M S
Leafbirds Chloropseidae
Cockatoos Cacatuidae
Yellow-throated Leafbird Chloropsis palawanensis P *
Red-vented Cockatoo Cacatua haematuropygia P
Greater Green Leafbird Chloropsis sonnerati B J M S
Parrots Psittacidae
Lesser Green Leafbird Chloropsis cyanopogon B M S **
Blue-crowned Hanging Parrot Loriculus galgulus B J M S ** Blue-winged Leafbird Chloropsis cochinchinensis B J M S
Yellow-throated Hanging Parrot Loriculus pusillus J
Bornean Leafbird Chloropsis kinabaluensis B *
Blue-rumped Parrot Psittinus cyanurus B M S
Sumatran Leafbird Chloropsis media S *
Blue-headed Racket-tail Prioniturus platenae P *
Orange-bellied Leafbird Chloropsis hardwickii M
Blue-naped Parrot Tanygnathus lucionensis B P
Blue-masked Leafbird Chloropsis venusta S *
Red-breasted Parakeet Psittacula alexandri J S
Flowerpeckers Dicaeidae
Long-tailed Parakeet Psittacula longicauda B M S **
Yellow-breasted Flowerpecker Prionochilus maculatus B M S **
Broadbills Eurylaimidae
Crimson-breasted Flowerpecker Prionochilus percussus B J M S **
Green Broadbill Calyptomena viridis B M S
Palawan Flowerpecker Prionochilus plateni P *
Hose's Broadbill Calyptomena hosii B *
Yellow-rumped Flowerpecker Prionochilus xanthopygius B *
Whitehead's Broadbill Calyptomena whiteheadi B *
Scarlet-breasted Flowerpecker Prionochilus thoracicus B M S
Black-and-red Broadbill Cymbirhynchus macrorhynchos B M S
Thick-billed Flowerpecker Dicaeum agile B J M S
Long-tailed Broadbill Psarisomus dalhousiae B M S
Striped Flowerpecker Dicaeum aeruginosum P
Silver-breasted Broadbill Serilophus lunatus M S
Brown-backed Flowerpecker Dicaeum everetti B M **
Banded Broadbill Eurylaimus javanicus B J M S
Yellow-vented Flowerpecker Dicaeum chrysorrheum B J M S
Black-and-yellow Broadbill Eurylaimus ochromalus B M S
Orange-bellied Flowerpecker Dicaeum trigonostigma B J M P S
Dusky Broadbill Corydon sumatranus B M S
Plain Flowerpecker Dicaeum minullum B J M S
Pittas Pittidae
Pygmy Flowerpecker Dicaeum pygmaeum P
Rusty-naped Pitta Hydrornis oatesi M
Blue-cheeked Flowerpecker Dicaeum maugei J
Schneider's Pitta Hydrornis schneideri S *
Black-sided Flowerpecker Dicaeum monticolum B *
Giant Pitta Hydrornis caeruleus B M S
Fire-breasted Flowerpecker Dicaeum ignipectus M S
Blue-headed Pitta Hydrornis baudii B *
Blood-breasted Flowerpecker Dicaeum sanguinolentum J
Javan Banded Pitta Hydrornis guajanus J *
Scarlet-backed Flowerpecker Dicaeum cruentatum B M S
Malayan Banded Pitta Hydrornis irena M S **
Scarlet-headed Flowerpecker Dicaeum trochileum J
Bornean Banded Pitta Hydrornis schwaneri B *
Sunbirds Nectariniidae
Red-bellied Pitta Erythropitta erythrogaster P
Ruby-cheeked Sunbird Chalcoparia singalensis B J M S
Blue-banded Pitta Erythropitta arquata B *
Plain Sunbird Anthreptes simplex B M S
Garnet Pitta Erythropitta granatina B M S **
Brown-throated Sunbird Anthreptes malacensis B J M P S
Graceful Pitta Erythropitta venusta S *
Red-throated Sunbird Anthreptes rhodolaemus B M S **
Black-crowned Pitta Erythropitta ussheri B *
Purple-naped Sunbird Hypogramma hypogrammicum B M S
Hooded Pitta Pitta sordida B J M P S
Purple-throated Sunbird Leptocoma sperata B P
Blue-winged Pitta Pitta moluccensis M
Van Hasselt's Sunbird Leptocoma brasiliana J M S
Mangrove Pitta Pitta megarhyncha M S
Copper-throated Sunbird Leptocoma calcostetha B J M P S
Australasian Warblers Acanthizidae
Olive-backed Sunbird Cinnyris jugularis B J M P S
Golden-bellied Gerygone Gerygone sulphurea B J M P S
Lovely Sunbird Aethopyga shelleyi P
Woodshrikes & Allies Tephrodornithidae
Handsome Sunbird Aethopyga bella P
Bar-winged Flycatcher-shrike Hemipus picatus B M S
White-flanked Sunbird Aethopyga eximia J *
Black-winged Flycatcher-shrike Hemipus hirundinaceus B J M S
**
Black-throated Sunbird Aethopyga saturata M
Large Woodshrike Tephrodornis virgatus B J M S
Crimson Sunbird Aethopyga siparaja B J M S
Rufous-winged Philentoma Philentoma pyrhoptera B M S **
Javan Sunbird Aethopyga mystacalis J *
Maroon-breasted Philentoma Philentoma velata B J M S ** Temminck's Sunbird Aethopyga temminckii B M S **
Bristlehead Pityriaseidae
Little Spiderhunter Arachnothera longirostra B J M S
Bristlehead Pityriasis gymnocephala B *
Pale Spiderhunter Arachnothera dilutior P *
Woodswallows and Allies Artamidae
Thick-billed Spiderhunter Arachnothera crassirostris B M S **
White-breasted Woodswallow Artamus leucorynchus B J M P S
Long-billed Spiderhunter Arachnothera robusta B J M S **
Ioras Aegithinidae
Spectacled Spiderhunter Arachnothera flavigaster B M S
Common Iora Aegithina tiphia B J M P S
Yellow-eared Spiderhunter Arachnothera chrysogenys B J M S **
Green Iora Aegithina viridissima B M S **
Grey-breasted Spiderhunter Arachnothera modesta M S
Great Iora Aegithina lafresnayei M
Streaky-breasted Spiderhunter Arachnothera affinis J *
Cuckooshrikes Campephagidae
Bornean Spiderhunter Arachnothera everetti B *
Large Cuckooshrike Coracina macei M
Streaked Spiderhunter Arachnothera magna M
Javan Cuckooshrike Coracina javensis J
Whitehead's Spiderhunter Arachnothera juliae B *
Sunda Cuckooshrike Coracina larvata B J S
Old World Sparrows Passeridae
Bar-bellied Cuckooshrike Coracina striata B J M P S
Plain-backed Sparrow Passer flaveolus M
Lesser Cuckooshrike Coracina fimbriata B J M S *
Eurasian Tree Sparrow Passer montanus J M P S
Pied Triller Lalage nigra B J M P S
Weavers Ploceidae
White-shouldered Triller Lalage sueurii J
Asian Golden Weaver Ploceus hypoxanthus J
Small Minivet Pericrocotus cinnamomeus J
Streaked Weaver Ploceus manyar J
Fiery Minivet Pericrocotus igneus B M P S ** Baya Weaver Ploceus philippinus J M S
Grey-chinned Minivet Pericrocotus solaris B M S
Waxbills Estrildidae
Sunda Minivet Pericrocotus miniatus J S
Red Avadavat Amandava amandava J
Scarlet Minivet Pericrocotus speciosus B J M S
Tawny-breasted Parrotfinch Erythrura hyperythra B J M S
Whistlers Pachycephalidae
Pin-tailed Parrotfinch Erythrura prasina B J M S
Mangrove Whistler Pachycephala cinerea B J M P S
White-rumped Munia Lonchura striata M S
White-vented Whistler Pachycephala homeyeri B
Javan Munia Lonchura leucogastroides J S **
Bornean Whistler Pachycephala hypoxantha B *
Dusky Munia Lonchura fuscans B *
Rusty-breasted Whistler Pachycephala fulvotincta J
Black-faced Munia Lonchura molucca J
Shrikes Laniidae
Scaly-breasted Munia Lonchura punctulata J M P S
Long-tailed Shrike Lanius schach B J M P S
White-bellied Munia Lonchura leucogastra B J M P S
**
Vireos & Greenlets Vireonidae
Black-headed Munia Lonchura malacca B
White-bellied Erpornis Erpornis zantholeuca B M S
White-capped Munia Lonchura ferruginosa J *
Pied Shrike-babbler Pteruthius flaviscapis J *
Chestnut Munia Lonchura atricapilla J M P S
Blyth's Shrike-babbler Pteruthius aeralatus M S
White-headed Munia Lonchura maja J M S
Black-eared Shrike-babbler Pteruthius melanotis M
Java Sparrow Lonchura oryzivora J *
Trilling Shrike-babbler Pteruthius aenobarbus J *
Wagtails & Pipits Motacillidae
Orioles Oriolidae
Paddyfield Pipit Anthus rufulus J M P S
Dark-throated Oriole Oriolus xanthonotus B J M P S
Finches Fringillidae
Black-naped Oriole Oriolus chinensis B J M P S
Brown Bullfinch Pyrrhula nipalensis M
Black-hooded Oriole Oriolus xanthornus B M S
Mountain Serin Chrysocorythus estherae J S
Black Oriole Oriolus hosii B *
Black-and-crimson Oriole Oriolus cruentus B J M S **
Gill F, Donsker D)2014) IOC World Bird List (v 4.4). URL http://www.worldbirdnames.org/
MacKinnon JR, Phillipps K (1999) A field guide to the birds of Borneo, Sumatra, Java and Bali. Oxford University Press
Phillipps Q, Phillipps K (2014) Phillipps' Field Guide to the Birds of Borneo, Third Edition. John Beaufoy, Oxford
Van Marle JG, Voous KH (1988) The birds of Sumatra. British Ornithologists' Union, Tring, Herts, United Kingdom
Wells DR (1999) The Birds of the Thai-Malay Peninsula. Vol. 1. Non-passeries. Academic Press, New York
Wells DR (2007) The Birds of the Thai-Malay Peninsula, Volume 2, Passerines. Christopher Helm, London, 800 pp
View publication statsView publication stats