Cockatoos Invade Indonesia,

Twice!
By J. Richard Wakefield
March 2004

Abstract

Cockatoos (Order Psittaciformes, family Cacatuidae) are distantly

related to the rest of the parrots, having separated some 20-30 million
years ago. They are very distinct in their appearance and habits.
Australia is their expected place of origin. They remained isolated as
that continent moved north due to plate tectonics. Once this plate
collided in the north into a complex series of other plates it formed the
Indonesian Archipelago. Cockatoos did not invade those islands until
about 5 million years ago, when there was enough closure for them to
make the flight across the sea. First into Papal New Guinea, and then
into the other islands. This invasion would have been into well-
established fauna and flora on these islands. Mitochondria DNA and
allozyme data produces a phylogenetic tree showing that there were
two separate invasions by the genus Cacatua. The order of invasion is
not known. One sub-genus, C Cacatua went westward into the smaller
islands and south to the Lesser Sunda Islands. They also invaded as
far north as the Philippines and east ward into the Solomon and
Bismarck islands. Some ambiguity persists on the actual order of the
chain of speciation events.

Wakefield, page 1
Introduction
Cockatoos belong to the Order Psittaciformes (parrots) which evolved as a group independent
of the rest of the Parrots. The Cockatoo family Cacatuidae is monophyletic (Christidis et al
1991).

They are a distinct group compared to the rest of their order by having a number of differences.
This includes the distinct head feather crest that can be raised and lowered, presence of a gall
bladder, powder down, features of the carotid arteries and skull. They also do not have Dyck-
texture in the feathers. In the other parrots this provides their feathers with the bright
colourations such as greens and yellows (Adams et al. 1984). Christidis et al. (1991) noted
that cockatoos have from 72 to 80 diploid number, where as the rest of the parrots have 60 to
72. Homberger (1996) also noted that the wood-ripping bill of cockatoos is distinctive
compared to the rest of the parrots, most likely being one of the major contributors to their
divergence.
Today they are restricted to Australia and the eastern portion of the Indonesian islands (
Moluccaian Islands, Tinambar Island, Sulawesi Island and the Lesser Sundra Islands.
), Papal New Guinea, Bismarck Islands, Solomon Islands and the Philippines.

Their origin is most likely Australia some time in on Gondwana some 20 to 30 million years
ago (Homberger 1991), with invasion into Indonesia, as we will see, much later. Though the
other parrots went on to colonize Africa and the New World, cockatoos were left stranded and
isolated on Australia upon its isolation from Antarctica until the collision with Indonesia
reduced the distance enough to cross into that region.

Interestingly, compared to the rest of the Parrots, cockatoos comprise of only 6 genera and
some 18 to 22 species. Cockatoos species range from abundant across the continent, to scarce
in some smaller regions, to small populations of several hundred on some of the Indonesian
islands.
They inhabit a wide range of habitats, such as forest, grasslands and tropical forest. Their diet
consists mostly of nuts, fruit, seeds and occasionally insect larva (which may have been their
original diet, [Homberger 1996]). They are considered very intelligent, capable of quick
learning, reasoning and have some problem solving abilities. Their beaks are well adapted for
ripping apart wood or seed cobs to get at the most edible parts (Homberger 1996). This has
lead to many of them to be considered crop pests ().

Generally, they are monogamous, and in the case of the Major Mitchell’s Cockatoo, require a
vary large range. Some, such as most of the Cacalua, are semi social living in flocks of 20 or
more, while others, such as the Palm Cockatoo, are solitary. Though no studies have yet
confirmed their longevity, in captivity they can live for up to 80 years. Breeding begins
around 2 years, with successful clutches after 5 years.

Wakefield, page 2
 Figure 1: Distance Wagner tree for Parrots of Australia rooted by Cockatoos. Though only three cockatoos, their
relationship different somewhat from Brown and Toft (1999), see below. From Christidis et al. 1991

All of this impacts an animal’s ability to colonize new areas as well as its ability to maintain a
viable population when their habitat changes, both of which occurred in the island areas of
Indonesia they invaded. New foods, competition from already entrenched species, and small
population sizes would hamper any founding populations (MacArthus and Wilson, 2001).
However, Cockatoo’s intelligence, curiosity and ability to exploit new foods certainly helped
in any new territory.

DNA phylogentic analysis has been used to determine the evolutionary history of the
cockatoos (Brown and Toft 1999, see below). More often than not, phyogentic trees produce
more than one cladogram. The island settings and the order of invasion should conform to the
phylogentic tree, or conversely can also determine past geological settings. Thus, the

Wakefield, page 3
geological setting of the Indonesian Islands would need to be explored to put any invasion into
context.

This paper will review the phylogentic relationships with that of the tectonic history of the
invasion areas. There are copious papers written on this subject for other fauna (for example
Ladiges et al 2003) where the special geological environment must be included in any study of
biogeography in Indonesia.

What can be shown is that, although the phylogeny may need some re-evalation or the
renaming of genera and species, it does for the most part support the geological setting. It
would appear that vicariant speciation and dispersal were the modes of speciation in the
Cacatua genus, the only genus to invade the north. In fact, one can infer that there were at
least two separate invasions from Australia by two different groups of cockatoos.

Indonesian Geology
There is only one small population cockatoos west of Wallace’s Line on a very small island,
hence our discussion on the geological setting will focus east of that line (See Appendix III).
The Indonesian archipelago is a mixture of island arc systems with raised and compressed sea
floor emerging as other islands

The tectonics of Indonesia is complex, with some 9 or more plates involved. None surpasses
the complexity of this system in Indonesia. In fact, much of the literature on other similar
systems often themselves reference pioneering work done in Indonesia (Charlton, 2000).
Though other such systems exist (Japan, Philippines, and the Caribbean islands), none are
thought to be more complex due to the plate interactions, or more tectonically active.

The Indonesian area, where cockatoos are found, was formed due to the Australian Plate,
considered independent of the Indian Plate (Sandiford et al, 2005), moving north colliding with
the Eurasian, Philippine, Pacific and a number of smaller plates. The relative motion of
Australia is approximately 7cm per year; one of the fastest plate motions measured anywhere
(Hall, 2001).

Papal New Guinea (PNG) is the logical stepping-stone for the movement of the cockatoos into
the islands, this will be shown later from the phylogeny. Thus it is important to understand
when and how it was formed to see if this confirms the phylogeny.

PNG is the upper most island of the Australian Plate. It was formed during the collision of the
Australian plate with a plate complex in the Pacific some 20 million years ago (Hall, 2001)

This is much too old for initial invasion by cockatoos as at that time the Australian continent
was too far south. Thus, the invasion of cockatoos in Indonesia must have happened long after
most of the islands were well established.

Wakefield, page 4
Figure 2: Basic Tectonics of the Indonesian and Australian area.

Figure 3: Tectonics of the eastern Indonesian area at 5 million years ago. Shaded land is current cockatoo ranges. From Hall,
R. (2001).

The Philippine islands, the first to form 30 milliion years ago, were much closer to PNG five
million years ago than they are today. The distance was an easy step for any cockatoo that
lived in PNG. The islands subsequently moved north and rotated, carrying with it the ancestor
of the species Cacatua haemaluropygia.

Wakefield, page 5
The Solomon and Bismarck islands to the west of PNG were formed from the collision of the
Pacific Plate with other smaller local plates. As these approached PNG, it became easier for
any PGN Cacutua to found those islands.

To the west, Ceram, Sulawesi and other smaller islands are the result of subduction induced
volcanic activity. The islands of Flores, Sumbawa and Lombok also are of island arc volcanic
origin.

One of the strangest chunks of land in the archipelago is Sumba Island. This island is the
southern most extension of the C. Sulphurea subspecies the Citron Cockatoo (C.s.
citrinocristata). This island is not volcanic, but actually a remnant of possible Sundaland 40-
50 ma consisting of continental crust that has been isolated for at least 30 million years
(Abdullah et al 2000, Hall 2001).

Timor island, east of Sumba Island, is also not an island arc origin, but from surfacing of
compressed crust during deformation.

Recent ice ages may have been a critical factor in the Cockatoo migrations. Though regardless
they could have migrated across the channel that separates the Gulf of Carpentaria from the
Gulf of Papau. Across this area the sea is very shallow, some 70 meters on average. During
the last ice age, sea levels were from 70 to 150 meters lower, exposing this and much of the
continental shelf as far west as Timor Island.

Life Takes Hold

Volcanic activity certainly played a major roll in the area. However, it is not likely that this
would have had much influence on Cockatoo invasion. The only influence of volcanism was
in the initial establishment of fauna and flora.

In August 26, 1883 one of the islands of Indonesia exploded. That eruption cleared the slate,
as it were, of all life on the local islands. Krakatau then became a scientist’s dream. It
allowed the systematic documentation of how life colonizes new islands.

This, then, can be used as model for how the islands in the whole archipelago became invaded
by life. New inhabitants could only come from two directions, west on the Java side and east
on the Sumatra side. The nearest undisturbed land was more than 40 kilometers away. Thus,
the only possible way life could have gotten to the new land was either by floating on the
currents, or airborne. Birds, it seemed, were very important in the dispersal of new plant life
on the island.

By Sept 1884, grass was noted for the first time on the remaining islands. By 1886 shore based
plants were seen growing on the island. By 1897, the interior was being inhabited, but what
was starting at that point was a clear division of fauna between the shoreline and the interior,
which has persisted today (Whittaker et al 1989).

Wakefield, page 6
Once birds and bats started to arrive, looking for food, or driven off course, various tree
species, such as figs, started to replace the grasses. The seeds being distributed by the
droppings (Whittaker and Jones 1994, Shilton et al 1999). With not much to eat, bird mortality
would have been high for the first arrivers.

However, within 50 years, the islands were well inhabited, though not completely. In fact, this
process is still occurring today, as new plants and animals arrive to take advantage of vacant
niches. Now, more than 50 different species of birds have been seen on the islands, of which
the vast majority are permanent residents.

In geological terms this is nothing. Extremely rapid re-colonization. This must have been
some of what would have happened as the whole chain of islands of Indonesia formed and
were colonized. Hence, the likelihood that cockatoos would have encountered a newly cleared
island is highly unlikely. Thus their invasion would have been on already well-established
land.

Extreme Biodiversity
In 1860 Alfred Russell Wallace noted that Indonesia was one of the most distinctive,
biologically, than any other place on the planet.

“The western and eastern islands of the archipelago belong to regions

more distinct and contrasted than any other of the great zoolological
divisions of the globe. (Quoted in Mayr, 1976)”

What was discovered is that the islands on the east side have fauna more closely related to
Australian fauna, and the western islands’ fauna almost entirely from the Asian main land.
The line where the two distinct ecosystems meet is now called the Wallace Line (see Appendix
III).

In fact, the number of bird species that have an Australian ancestry decrease in numbers as one
moves westward across the archipelago. Cockatoos also seemed to have followed the same
pattern of dispersal.

Cockatoo Phylogeny
Brown and Toft (1999) performed mDNA cladistic analysis on most of the Cockatoo species to
get a phylogenic relationship. They utilized what Adams et al (1984) had already done with
allozyme analysis on 10 species of cockatoos. The Brown and Toft’s group also excluded
some of the species and subspecies (see Appendix I). They were only interested in the Generic
level of relationships. Out Group species, which they included, were the Rock Dove, Common
Canary and the Japanese Quail, noting the justifications for doing so.

They also mapped into this the geographic location, giving each range a value of either present
or not present in the ancestral range.

Wakefield, page 7
Wakefield, page 8
Alloyme data came from Adams et al (1984), but Brown and Toft did all the mDNA extraction
and analysis.

Their results were processed using a number of methods, which produced a number of equally
parsimonious trees. They then took the 12 resultant trees and created a majority-rule consensus
tree (see Figure 4)

For our purposes we need to see the Cacatua genre only. In fact, their Strict-Consensus tree of
just their own mDNA analysis shows very nicely this phylogeny, and as we will see, fits well
into the proposed invasion sequence.

Brown and Toft (1999) surmised from their data and trees that two distinct groups of cockatoos
formed the Cacatua clade, which
in fact has been suggested for
some time from morphological
data (Adams et al 1984).

They also discovered that the

Major Mitchel’s Cockatoo (C.
leadbeateri) does not belong in
the genus Cacatua even though
some of the morphology is close
(and hence shows as a sister
group to the Cacatua).

Figure 5: Cacatua Phylogeny based on mDNA analysis. Two clear groups They also confirmed that
of sub-genera stand out, which fits well with their two distinct invasions of cockatiels are part, though
Indonesian islands. From Brown and Toft (1999) distantly, of the Cockatoo family.

Cockatoo Invasion
Thus with all of the above understood, it is likely that a reconstruction of Cockatoo invasion
sequence can be attempted. However, missing from above is any attempt of the age at which
each speciation (node) in the Cacatua clade took place. Brown and Toft (1999) did not
produce a Distance Wagner tree (though one can be done from the data they published, it is
beyond the scope of this paper), and thus mapping to the tectonic events is problematic.

What is clear is with the C. (Licmetis) clade some ancestor of C. Sanguinea invaded PNG from
Australia, splitting then to the Solomons (C. Ducorpsii), Philippines ( C. haemaluropygia) and
Tanimbar Island (C. goffini).

Brown and Toft (1999) mDNA Strict Consensus tree (figure 5) shows those three species C.
sanguinea, gofini and ducorpsii are triplets. Is it possible that these last two split
simultaneously, one going off in each direction (see figure 6)

Wakefield, page 9
 C. haemaluropygia was able to
invade the Philippines because that
set of islands was much closer to
PNG some time around 5ma. This
shows clearly in that this species is
way at the bottom of the Licmetis
sub-genera showing a very early
split. But without some indication as
to when this splitting (speciation)
took place there is not much
correlation as to when the islands
west of PNG were invaded.

However, the phylogeny of the

subgenus C. (Cacatua) has a little
ambiguous understanding of what
happened due to the different
cladograms produce by Brown and
Toft (see figures 7A and 7B). The
invasion, or what’s left of it, follows
a set of islands to the west of PNG,
with one heading to the east onto the
Bismarck Archipelago (C.
ophthamica).

Brown and Toft’s tree, which

includes both the allozyme and
mDNA data, seems to imply that
some ancestor to C. molucansis, C.
alba, C. ophthamica and C.
sulphurea invaded PNG. Remnants
of that ancestor may have lived on in
the Triton Cockatoo, a sub-species of
C. galaria. From that ancestor the
invasion apprears to have been
westward twice, if the tree in figure 6
is a proper indication of the
phylogeny. C. sulphurea is the
furthest west from PNG, but is the closest to this ancestor, with C. alba and C. moluccansis
evolving later. This seems unlikely.

However, if their mDNA only tree in Figure 5 is more correct, we have a better indication of
what might have happened because we have a triplet between C sulphurea, C. ophthamica and
the ancestor to C. alba and C. moluccansis. C. ophthamica went eastward around the same
time as the ancestor to C sulphurea + C. alba + C. moluccansis went westward. Then this
ancestor then went on to invade the Sulawesi Islands where C sulphurea evolved, but leaving

Wakefield, page 10
Wakefield, page 11
Wakefield, page 12
behind a population on the series of Moluccian islands which then separated into C. alba and
C. moluccansis (Figure 7B).

Brown and Toft (1999), claim that the islands on which C. moluccansi is located (Moluccian)
and the Bismarck islands (C. ophthamica) were once tectonically joined, hence, they argued,
supports the closeness of the two species according to their cladogram in Figure 4. These two
species then became separated geographically when PNG formed. This is somewhat hard to
accept and no geological evidence was found to support their hypothesis. Unless more data is
gathered and incorporated in this phylogeny, such questions remain a mystery.

The subspecies of C. sulphurea then invaded the Lesser Sunda Islands south of Sulawesi Island
(see figure 7). C.s. citrinocristata is found only on Sumba island while C.s. abbotti is found
only on Solombo Besar Island, west of Wallace’s Line, which should make both of them the
last in the phylogeny and the newest speciation. C.s. parvula on Timor and Samao Island, C. s.
occidentalis is found on Lombok, Sumbawa and Flores, and Noesa Penida Island, and C. s.
djampeana on the islands of Islands of Alor, Pantar, Djampea, Kalao, Kalao tua, Madu and
Kaju adi, and Tukangbesi Islands

Figure 7. Map of invasion of the subspecies of the Cacatua Sulphurea into the Lesser Sunda
Islands. Actual times of invasions and eventual permanent inhabitation are unknown.

Wakefield, page 13
Cockatoo invasion would follow the equilibrium theory of island biogeography (MacArthur
and Wilson 2001) which basically states:
The rate of species successful invasion decreases with the number of
resident species on an island and the extinction rate increases with the
number of resident species.

Since it appears cockatoos invaded already established areas, they would have been subject to
this kind of pressure. Thus, it is very likely that some of these dispersal populations did not
make it, and that skews the expected biogeographic reconstruction.

Conclusion

Not all of the phylogeny of cockatoos has been worked out completely. The uniqueness of the
subspecies, and the specific dispersals onto islands of species not in the Brown and Toft study
would miss specific details. However, their goal was not to go beyond the genera level.

The relatedness of each of the species and subspecies could solve a number of remaining
questions. Without the rest of the species and subspecies included, it is difficult to get a good
understanding as to the chain of events that lead to the invasion of Indonesia. Working out
phylogenetically the subspecies of C. sulphurea, for example, could tell us the order of
invasion, and approximately when each split took place.

Missing also from the current mDNA of Brown and Toft’s analysis was a Distance Wagner
tree. This could provide a timeline as to when some of these speciation events took place.

It could also answer the similarities between the Blue-Eyes and the Moluccan similarity other
than the one proposed by Brown and Toft that does not fit well with the known tectonic events.

It also may answer as to why the White Cockatoo lost its crest display colours with respect to
the rest of the subgenus it belongs to (it also raises those crest feathers quite differently than the
Moluccian or Sulphur Crested, personal observation).

Generally, however, the basic understanding of how the invasion took place matches the
tectonic chain of events. There were two distinct invasions, one of the Cacatua subgenus and
the other of the Licmetis subgenus. A Distance Wagner tree may resolve which subgenus was
first into Papal New Guinea, or if it was simultaneous invasion.

It appears, though not confirmed, that cockatoos invaded the Indonesian Archipelago very late
in their history, less than 5ma, thus at that time most of the islands were faunally well
established. Cockatoos would have been a new invasion as far as the other species present
were concerned. Cockatoos certainly have the tools (flying ability, their strong beak, wide
food range, and a nimble zygodactyle feet) to take on any new environments.

One of the more interesting questions still to be answered is why the Cacatua are all white?
Very conspicuous indeed. What selection pressure changed them from black the prominent
colour of the other cockatoos?

Wakefield, page 14
On a side note, invasions into new territories mean higher selection pressures, coupled with
smaller less stable populations, means they are more prone to extinction. Though the pet trade
of this birds has had its toll, certainly now the biggest threat to the Cacatua species is loss of
habitat due to over logging. Efforts are underway to help save these birds from extinction,
though some populations may have already gone.

Wakefield, page 15
 Appendix I
Cocaktoo Common and Species names. Bolded use in study by Brown and Toft (1999)
Genera Species Common Range
Cacatua alba White (Umbrella) Cockatoo Aru Islands
ducorpsii Ducorps Cockatoo Solomon Islands
galarira galerita Sulphur Crested Cockatoo North-east coast Australia
galarira triton Triton Cockatoo Papua New Guinea
goffini Goffins Cockatoo Tanimbar Is. Kai Is.
haemaluropygia Philippine Cockatoo Philippines
leadbeateri Major Mitchells Cockatoo Central and Southern Australia
moluccansis Moluccan Cockatoo Southern Moluccas: Seram, Saparua,
Haruku
ophthamica Blue-eyed cockatoo New Britain, New Ireland (Bismarck)
pastinator Western Corella
sanguinea Little Corella Throughout Australia
sulphurea abbotti Medium Sulphur Crested Solombo Besar Island
Cockatoo
sulphurea citrinocristata Citron Crested Cockatoo Sumba Is.
sulphurea djampeana Islands of Alor, Pantar, Djampea, Kalao,
Kalao tua, Madu and Kaju adi, and in the
Tukangbesi Islands
sulphurea occidentalis Lombok, Sumbawa and Flores, Noesa
Penida Island
sulphurea parvula Timor and Samao Island
sulphurea sulphurea Lesser Sulphur (Yellow) Celebes and adjacent Buton Island,
Crested Cockatoo introduced to Singapore
tenuirostis Slender Billed Corella
Eolophus rosaicapittus Galah Cockatoo
Callocephaion fimbriatum Gang-gang cockatoo Extreme south-west Australia
Calyptorhynchus lathami Glossy Cockatoo South-Eastern Australia
baudinii Long-billed Black-Cockatoo Australia
magnificus Red-Tailed black cockatoo Australia
funereus funereus Yellow-tailed black cockatoo Australia
funereus baudinii White-tailed black cockatoo Australia
latirostris Slender-billed Black-Cockatoo Australia
banksii Red-tailed Black-Cockatoo Australia
Nymphlous hollandicus Australia
Proboscigar aferrimus aterrimus Palm Cockatoo Cape of York Peninsula in Northern
Australia
aferrimus goliath Goliath Palm Cockatoo New Guinea
aferrimus stenolophus Very North-west tip of New Guinea

Wakefield, page 16
 Appendix II, Identification

Licmetis clade
Philippine Cockatoo Goffins Cockatoo

Little Corella Ducorps Cockatoo

Western Corella Slender Billed Corella

Wakefield, page 17
 Cacatua Clade
Sulphur Crested Cockatoo Triton Cockatoo

Moluccan Cockatoo White (Umbrella) Cockatoo

Blue-eyed cockatoo Lesser Sulphur (Yellow) Crested Cockatoo

Wakefield, page 18
Citron Crested Cockatoo Medium Sulphur Crested Cockatoo

Non-Cacatua Clades
Major Mitchells Cockatoo Galah Cockatoo

Wakefield, page 19
 Gang-gang cockatoo Glossy Cockatoo

Long-billed Black-Cockatoo Red-Tailed black cockatoo

Yellow-tailed black cockatoo White-tailed black cockatoo

Wakefield, page 20
Palm Cockatoo

Wakefield, page 21
 Appendix III
Geography of Indonesian Archipelago

Wakefield, page 22
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