Journal of Biogeography (J. Biogeogr.) (2010) 37, 1767–1781

ORIGINAL
ARTICLE

Biogeographical history of cuckoo-shrikes(Aves: Passeriformes): transoceanic
colonization of Africa from
Australo-Papua
Knud A. Jønsson1,2*, Rauri C. K. Bowie2, Johan A. A. Nylander3,
Les Christidis4,5, Janette A. Norman5,6 and Jon Fjeldså11Vertebrate Department, Zoological Museum,
University of Copenhagen, Universitetsparken
15, DK-2100 Copenhagen Ø, Denmark,2
Museum of Vertebrate Zoology andDepartment of Integrative Biology, 3101 Valley
Life Science Building, University of California,
Berkeley, CA 94720-3160, USA, 3Departmentof Botany, Stockholm University, S-106 91
Stockholm, Sweden, 4National Marine ScienceCentre, Southern Cross University, Coffs
Harbour, NSW 2450, Australia, 5Departmentof Genetics, University of Melbourne, Parkville,
Vic. 3052, Australia, 6Sciences Department,
Museum Victoria, GPO Box 666, Melbourne,
Vic. 3001, Australia

ABSTRACTAim Cuckoo-shrikes and allies (Campephagidae) form a radiation of birds
widely distributed in the Indo-Pacific and Africa. Recent studies on the group
have been hampered by poor taxon sampling, causing inferences aboutsystematics and biogeography to be rather speculative. With improved taxon
sampling and analyses within an explicit spatiotemporal framework, we elucidate
biogeographical patterns of dispersal and diversification within this diverse cladeof passerine birds.
Location Africa, Asia, Australo-Papua, the Pacific, the Philippines and Wallacea.
Methods We use model-based phylogenetic methods (MrBayes and garli) toconstruct a phylogenetic hypothesis of the core Campephagidae (Campephagidaewith the exclusion of Pericrocotus). The phylogeny is used to assess thebiogeographical history of the group with a newly developed Bayesian
approach to dispersal–vicariance analysis (Bayes-diva). We also made use of apartitioned beast analysis, with several calibration points taken from island ages,
passerine mitochondrial substitution rates and secondary calibration points forpasserine birds, to assess the timing of diversification and dispersal.
Results We present a robust molecular phylogeny that includes all genera
and 84% of the species within the core Campephagidae. Furthermore, weestimate divergence dates and ancestral area relationships. We demonstratethat Campephagidae originated in Australo-Papua with a single lineage(Pericrocotus) dispersing to Asia early. Later, there was further extensive
transoceanic dispersal from Australo-Papua to Africa involving lineages within
the core Campephagidae radiation.
Main conclusions The phylogenetic relationships, along with the results of theancestral area analysis and the timing of dispersal events, support a transoceanic
dispersal scenario from Australo-Papua to Africa by the core Campephagidae.
The sister group to core Campephagidae, Pericrocotus, dispersed to mainland Asia
in the late Oligocene. Asia remained uncolonized by the core Campephagidae
until the Pliocene. Transoceanic dispersal is by no means an unknown
phenomenon, but our results represent a convincing case of colonization
over a significant water gap of thousands of kilometres from Australo-Papua
to Africa.

*Correspondence: Knud A. Jønsson, Vertebrate
Department, Zoological Museum, University of
Copenhagen, Universitetsparken 15, DK-2100
Copenhagen Ø, Denmark.
E-mail: kajonsson@snm.ku.dkª 2010 Blackwell Publishing Ltd

Keywords
Africa, Australia, biogeography, birds, Campephagidae, cuckoo-shrikes, IndoPacific, Passeriformes, phylogeny, transoceanic dispersal.

www.blackwellpublishing.com/jbi
doi:10.1111/j.1365-2699.2010.02328.x

1767

K. A. Jønsson et al.
INTRODUCTION
Recent advances in our understanding of biological dynamics
within islands and archipelagos, and between archipelagos andcontinents, have demonstrated a greater complexity thanthought by earlier biogeographers (reviewed by Whittaker &Fernández-Palacios, 2007; Bellemain & Ricklefs, 2008; Emerson & Gillespie, 2008; Gillespie et al., 2008). In particular, theincreasing significance of reverse colonizations, whereby island
systems are sources for continental biodiversity, has challenged
a fundamental biogeographical paradigm (Filardi & Moyle,
2005; Bellemain & Ricklefs, 2008).
A second paradigm within biogeography that is receiving
renewed attention is the determination of the relative importance of dispersal and vicariance in shaping distribution
patterns of organisms across the globe (de Queiroz, 2005;
Heaney, 2007). Dispersal and vicariance are often considered
competing hypotheses when explaining present distribution
patterns (de Queiroz, 2005). For example, disjunctly distributed sister-species can be explained either by dispersal or bypostulating that an ancestral taxon was historically widespread,and that the present distribution pattern is a consequence ofthe extinction of populations from the intervening area. The
advent of molecular-based phylogenies with corresponding
divergence time estimates, however, has started to provide data
sets with which one can explicitly test the underlying historicalbiogeographical pattern (e.g. Sanmartı́n & Ronquist, 2004;
Brown et al., 2006; Nylander et al., 2008b; Voelker et al.,2009).
The avian family Campephagidae (cuckoo-shrikes andallies) is distributed throughout Africa, Asia and Australia, as
well as within the archipelagos of both the Indian and Pacific
Oceans. Consequently, gaining an understanding of the origin,pattern and timing of dispersal that have led to the present
cuckoo-shrike radiation is likely to be integral to interpreting
broader regional biogeographical patterns and processes acrosscontinents and archipelagos.
The origin of passerine birds (Passeriformes) has been
demonstrated to be within the Gondwanan supercontinentand ranges back to the Cretaceous/Tertiary (K/T) boundary(Barker et al., 2002, 2004; Ericson et al., 2002). Two major
lineages within Passeriformes are recognized: the suboscines(Tyranni), which occur primarily in South America; and theoscines (Passeri), with a putative Australian origin. The basal
lineages within oscines all occur in Australia, thus it is likelythat Australia is the place of origin for this highly diverse
radiation (Christidis, 1991; Barker et al., 2002; Edwards &Boles, 2002; Ericson et al., 2002).
One of the major avian biogeographical questions in recent
years has been how oscines dispersed out of Australia. One
sub-group, the Passerida, which has been very successful interms of diversity (c. 3500 species), is thought to haveoriginated in Africa (Fuchs et al., 2006; Jønsson & Fjeldså,2006; Johansson et al., 2008). For this hypothesis to be correct,
it would require transoceanic dispersal from Australia to Africa
of an early Passerida ancestor, a scenario perhaps more
1768

parsimonious than assuming a partial extinction of a once
widespread common ancestor/s.
Another large radiation within the Passeri is the core
Corvoidea (c. 750 species, sensu Monroe & Sibley, 1993). The
Campephagidae is the most species-rich family of birds bothwithin the core Corvoidea complex, and in the Indo-Pacificregion. The core Campephagidae, which excludes Pericrocrotus(minivets), but includes the cuckoo-shrikes and trillers
belonging to the genera Coracina, Lalage, Campochaera,
Campephaga and Lobotos, numbers 70 species, 57 of which
are distributed from India (including Sri Lanka) in the west toAustralia and the Pacific Islands in the east. A further 11
species occur in tropical Africa (including Madagascar and theComoros), and two species occur on the Indian Ocean islandsof Mauritius and Réunion. The systematic relationships within
this group are poorly studied, and consequently inferences
made on the biogeographical history are highly speculative.
It has been argued that the cradle of campephagid diversification was within the Australo-Papuan region (Jønsson
et al., 2008). However, the alternative hypothesis – that Asia
could be the area of origin from where colonization of Africa
and Australo-Papua could be explained by one radiation goingwest to Africa and another going east to Australo-Papua –could not be excluded. Recent work by Fuchs et al. (2007) andJønsson et al. (2008) has suggested close affinities between
African and Australian/Wallacean cuckoo-shrike species, leading the authors to speculate that dispersal directly between thetwo continents was a likely scenario. These studies, however,
suffered from relatively poor taxon sampling hampering amore substantiated biogeographical investigation of the group.
In the present study we construct a densely sampled molecularphylogeny, which we use to assess several a priori hypotheses inorder to gain a better understanding of the biogeographicalhistory of cuckoo-shrikes. If cuckoo-shrikes originated in Asia,
we would expect Asian taxa to dominate the basal clade(s)within the phylogeny. Conversely, if the origin was in AustraloPapua, then these taxa would be expected to dominate the basal
part of the tree. If dispersal took place from Australia to Africa
via Asia, as the terranes of the Australian and Asian plates
collided some 20 Ma (Hall, 1998, 2002; Holloway, 1998), wewould expect to find multiple dispersal events reflecting theclose proximity of the many larger and smaller islands in theIndonesian and Philippine archipelagos, which would have
served as stepping stones between Australia and mainland Asia.
This kind of dispersal pattern has been demonstrated for othergroups of birds, for example, the Pachycephalidae (Jønsson
et al., 2010a). These Asian colonizers would, in turn, be expected
to be the founders of African lineages, and we would expect tofind African lineages at distal parts of the tree.
MATERIALS AND METHODS
Taxon sampling and laboratory procedures
Recent studies of higher-level relationships within the Campephagidae have demonstrated that the genera Lobotos,
Journal of Biogeography 37, 1767–1781ª 2010 Blackwell Publishing Ltd

Phylogeny and biogeography of cuckoo-shrikes
Campephaga, Campochaera and Lalage are nested within the
Coracina complex. Pericrocotus, on the other hand, is themonophyletic sister-group (Fuchs et al., 2007; Jønsson et al.,2008). Sister to the Campephagidae (comprising Coracina andPericrocotus) is an African assemblage including the families
Platysteiridae and Malaconotidae (Fuchs et al., 2007; Jønsson
et al., 2008), which are intermingled with some Australasianspecies (Norman et al., 2009). Although the primary emphasis
in our current paper is on the Coracina complex, we included
several taxa from known sister groups (e.g. Pericrocotus andMalaconotidae) in order to address biogeographical questions
appropriately. With this in mind, we have sampled 59 of the
70 (84%) species included within the core Campephagidae(Taylor, 2005).
Fresh tissue (blood, liver or muscle) was extracted using the
Qiagen DNeasy Extraction kit (Qiagen, Valencia, CA, USA),
following the manufacturer’s protocol. Three nuclear generegions: myoglobin intron-2 (Myo2) (chromosome 1), ornithine decarboxylase (ODC) intron-6 to intron-7 (chromosome3), and glyceraldehyde-3-phosphodehydrogenase (GAPDH)
intron-11 (chromosome 1); and two mitochondrial markers:
NADH dehydrogenase subunit 2 (ND2) and subunit 3 (ND3),
were sequenced and used to estimate phylogenetic relationships. Primer pairs used for amplification were: ND2: Lmet
(Hackett, 1996)/H6312 (Cicero & Johnson, 2001); ND3: ND3L10755/ND3-H11151 (Chesser, 1999); myoglobin intron-2:
Myo2 (Slade et al., 1993)/Myo-cora2R (Jønsson et al., 2008)
and Myo-coraF1 (Jønsson et al., 2008)/Myo3F (Heslewood
et al., 1998); ODC: OD6/OD8 (Allen & Omland, 2003) andG3P13/G3P14b (Fjeldså et al., 2003).
The thermocycling conditions included a hot start at 95 C
for 5 min, followed by 32 cycles at 95 C for 40 s, 54–63 C for
40 s, and 72 C for 60 s, and was completed by a final
extension at 72 C for 8 min. One microlitre of the amplification products was electrophoresed on a 1.5% agarose gel andrevealed under UV light with ethidium bromide to check forcorrect fragment size and to control for the specificity of theamplifications. Polymerase chain reaction (PCR) products
were purified using ExoSap enzymes (exonuclease and shrimp
alkaline phosphatase). Purified PCR products were cyclesequenced using the Big Dye terminator chemistry (ABI;
Applied Biosystems, Foster City, CA, USA) in both directions
with the same primers as were used for PCR amplifications,
except for G3P13, which was replaced by G3PintL1 (Fjeldså
et al., 2003), and run on an automated ABI 3100 DNA
sequencer.
Corresponding laboratory procedures for study skins aredetailed by Irestedt et al. (2006). Additional internal primers
for study skins are specified by Jønsson et al. (2008) for
myoglobin and GAPDH, and by Irestedt et al. (2006) for ODC.
In addition, two new primers were specifically designed for thisstudy, ND2cor778R: ATGATGAGTCAT TTTGGGAGGAA
and ND2cor795F: CAGGTTTCCTCCCAAAGTGACTC.
Sequences were assembled with SeqMan II (DNASTAR Inc.,
Madison, WI, USA). Positions where the nucleotide could not
be determined with certainty were coded with the appropriate
Journal of Biogeography 37, 1767–1781ª 2010 Blackwell Publishing Ltd

IUPAC code. GenBank accession numbers are provided inTable 1.
Alignment and phylogenetic analyses
Alignment was performed using MegAlign (DNASTAR Inc.)
with some minor manual adjustments. The concatenatedalignment consisted of 3069 bp. Coding genes (ND2 and ND3)
were checked for the presence of stop codons or insertion/
deletion events that would have disrupted the reading frame.
We used Bayesian inference (e.g. Huelsenbeck et al., 2001;
Holder & Lewis, 2003), as implemented in MrBayes 3.1.2(Huelsenbeck & Ronquist, 2003; Ronquist & Huelsenbeck,2003) to estimate phylogenetic relationships. The mostappropriate substitution models were determined with
MrModeltest 2.0 (Nylander, 2004), using the Akaike information criterion (Akaike, 1973; Posada & Buckley, 2004).
Bayesian analyses for the concatenated data set were performed
allowing the different parameters (base frequencies, rate matrix
or transition/transversion ratio, shape parameter, proportionof invariable sites) to vary between the seven partitions(GAPDH, ODC, Myo2, 1st, 2nd, 3rd codon positions for
mtDNA, and tRNA), i.e. mixed-models analyses (Ronquist &Huelsenbeck, 2003; Nylander et al., 2004). In all MrBayes
analyses, the Markov chain Monte Carlo (MCMC) was run
using Metropolis-coupling, with one cold and three heatedchains, for 5 million to 15 million iterations with trees sampled
every 1000 iterations. The number of iterations discarded
before the posterior probabilities were calculated (i.e. thelength of the ‘burn-in’ period) was estimated graphically using
AWTY (Wilgenbusch et al., 2004; Nylander et al., 2008a) bymonitoring the change in cumulative split frequencies. Two
independent runs initiated from random starting trees were
performed for each data set, and the log-likelihood values andposterior probabilities for splits and model parameters werechecked to ascertain that the chains had reached apparent
stationarity.
Maximum-likelihood analyses were performed using garli
0.95 (Zwickl, 2006). Five independent analyses were performed(20 million generations for the combined analysis, 15 million
generations for ND2, and 10 million generations for the
nuclear partitions). Nodal support was evaluated with 500
nonparametric bootstrap pseudoreplications.
Establishing ancestral areas
In order to elucidate ancestral patterns we used the newly
developed ‘Bayes-diva’ approach (Nylander et al., 2008b). In astandard dispersal–vicariance analysis (Ronquist, 1997) as
implemented in the software diva (Ronquist, 1996, 2001),
ancestral areas are optimized onto internal nodes of aphylogeny by minimizing the number of dispersal andextinction events required to explain the terminal distributions(Ronquist, 2003). The basic assumption (null model) is
allopatric speciation in response to vicariance, but diva also
considers dispersal and extinction in the shaping of current
1769

K. A. Jønsson et al.
Table 1 List of taxa included in the study.
Taxon name

Voucher number

Origin

GAPDH

ODC

Myo

Aleadryas rufinucha
Artamus cyanopterus
Batis poensis
Campephaga flava
Campephaga petiti
Campochaera sloetii
Coracina abbotti
Coracina analis
Coracina atriceps
Coracina azurea
Coracina bicolor
Coracina boyeri
Coracina caeruleogrisea
Coracina caesia
Coracina caesia
Coracina caledonica
Coracina ceramensis
Coracina cinerea
Coracina coerulescens
Coracina dispar
Coracina dohertyi
Coracina fimbriata
Coracina graueri
Coracina holopolia
Coracina incerta
Coracina ingens
Coracina larvata normani
Coracina leucopygia
Coracina lineata lineata
Coracina lineata sublineata
Coracina longicauda
Coracina macei larvivora
Coracina macei larvivora
Coracina maxima
Coracina mcgregori
Coracina melanoptera
Coracina melaschistos
Coracina mindanensis everetti
Coracina montana
Coracina morio
Coracina newtoni
Coracina novaehollandiae
Coracina ostenta
Coracina papuensis artamoides
Coracina papuensis sclaterii
Coracina pectoralis
Coracina polioptera
Coracina salomonis
Coracina schistacea
Coracina striata guillemardi
Coracina striata kochii
Coracina sula
Coracina temminckii
Coracina tenuirostris admiralitatis
Coracina tenuirostris heinrothi
Coracina tenuirostris matthiae
Coracina tenuirostris remota

NRM 543658
ZMUC 135911
MNHN CG 1998-783
RB 613
FMNH 384855
NRM 569328
NRM 569468
AMNH 337784
BMNH 1910.12.28.182
ANSP 11901
RMNH.AVES 61247
ANWC 27158
ANWC 24169
ZMUC 123521
ZMUC 134772
FMNH 268458
AMNH 562444
FMNH 352837
USNM B3671
AMNH 562297
RMNH.AVES 85029
RMNH.AVES 66951
RG 79-33-A-2
AMNH MKL80
ANWC 27044
AMNH 334884
BMNH 1935.10.22.303
RMNH.AVES 123485
ANWC 39960
ZMUC 95267
ANWC 26970
MNHN CG 1989-68
ZMUC 95260
ANWC 48004
FMNH 392259
FMNH 233770
MNHN 6-69
ZMUC 95262
ANWC 24172
RMNH.AVES 61242
SEOR 001
ANSP 10622
AMNH 459820
ANWC 39813
ZMUC 95265
MNHN CG 1979-1352
MNHN CG 1989-69
ZMUC 139341
AMNH 561078
ZMUC 95261
ZMUC 95258
AMNH 562441
NRM 569324
ZMUC 95268
ZMUC 95264
ZMUC 95266
ZMUC 102478

New Guinea
Australia
Cameroon
Kenya
UgandaNew Guinea
Sulawesi
New Caledonia
Ceram
Equatorial Guinea
Sulawesi
New Guinea
New Guinea
Tanzania
AfricaLoyalty Isl
Ceram
Madagascar
Philippines
Little Key
FloresSumatraCongo
Solomon Islands
New Guinea
Manus
BorneoSulawesi
Australia
New Ireland
New Guinea
ThailandThailandAustralia
Philippines
India
Laos
Tawi Tawi
New Guinea
Sulawesi
Réunion
Australia
Philippines
Australia
New Britain
South Africa
Thailand
MakiraSula
Tawi Tawi
Mindanao
Sula
Sulawesi
Manus Island
New Britain
MussauNew Hanover

EU273375
DQ406661
DQ406665
DQ406639
EF052795
EU380459
EU380460
HM002899
EU272091
EF052796
HM002900
HM002901
HM002902
EF052797

EU273355

EU273395
DQ406636
AY529907
EF052822
EF052823
EU380489
EU380491

1770

EU272120
EU380410
EU380411
EU380412
EU380415
HM002997
EU273118
EU380416
HM002998
HM002999
HM003000

EU272102
EF052824
HM002926
HM002927
HM002928
EF052825

HM002903

HM003001
HM003002

HM002929

EF052800
EF052759

EU380417
EU380418

EF052827
EF052770

HM002904
HM002905
HM002906
EF052801
HM002907

HM003003
HM003004
HM003005
HM003006

HM002930
HM002931
HM002932
EF052828
HM002933

EU380461
HM002908
HM002909
EU380462
HM002910

EU380419
HM003007
HM003008
EU380420
HM003009

EU380492
HM002934
HM002935
EU380493
HM002936

EU380463
HM002911
EF052805
HM002912
EF052807
EU380464
HM002913
HM002914
HM002915
EF052808
HM002916
HM002917
EU380465
EF052810
EF052811
HM002918

EU380421
HM003010
EU380422
HM003011
EU380423
EU380424
HM003012
HM003013
HM003014
EU380425
HM003015
HM003016
EU380426

EU380494
HM002937
EF052831
HM002938
AY529913
EU380495
HM002939
HM002940
HM002941
EF052834

HM003017

HM002942
EU380496
EF052836
EF052837
HM002943

EU380466
EU380467

EU380427
EU380428

EU380497
EU380498

EU380468
EU380469
EU380470
EU380471
HM002919

EU380429
EU380430
EU380431
EU380432
HM003018

EU380499
EU380500
EU380501
EU380502
HM002944

ND2

DQ096728
AY529941
AY529944
EF052771
HM002951
HM002952
HM002953
EF052772
HM002954
HM002955
HM002956
EF052773
HM002957
HM002958
EF052685
EF052697
HM002959
HM002960
HM002961
HM002962
EF052775
HM002963
HM002964
HM002965
HM002966
HM002967
EF052777
HM002968
EF052686
HM002969
AY529948
HM002970
HM002971
HM002972
HM002973
EF052779
HM002974
HM002975
HM002976
EF052781
EF052782
HM002977
HM002978HM002979
HM002980
HM002981
HM002982
HM002983
HM002984
HM002985
HM002986

Journal of Biogeography 37, 1767–1781ª 2010 Blackwell Publishing Ltd

Phylogeny and biogeography of cuckoo-shrikes
Table 1 Continued
Taxon name

Voucher number

Origin

GAPDH

ODC

Myo

ND2

Coracina tenuirostris tenuirostris
Coracina typica
Coracina welchmani
Daphoenositta chrysoptera
Eopsaltria australis
Gymnorhina tibicen
Lalage atrovirens
Lalage leucomela
Lalage leucopyga
Lalage leucopygialis
Lalage maculosa
Lalage melanoleuca minor
Lalage nigra
Lalage sharpei
Lalage sueurii
Lalage tricolor
Lobotos oriolinus
Machaerirhynchus flaviventer
Malaconotus blanchoti
Oreoica gutturalis
Oriolus oriolus
Ornorectes cristatus
Orthonyx temminckii
Pachycephala pectoralis
Peltops blainvillii
Pericrocotus divaricatus
Pericrocotus erythropygius
Platysteira cyanea
Prionops retzii
Rhagologus leucostigma
Telophorus sulfureopectus
Outgroup
Menura novaehollandiae

ANWC 41992
ZMUC 141101
UWBM 60241
MV 1311
MV 1390
MV AC78
RMNH.AVES 84504
UWBM 57519
ZMUC 139386
BMNH 1934.10.21137
AMNH 206135
ZMUC 95259
FMNH 344979
AMNH 206892
RMNH.AVES 65417
UWBM 57508
RG 126479
ANWC 39520
ZMUC 116824
SAM B.39217
MCSNC 1415
ANWC 26733
MVB 831
MV 3477
ANWC 26492
UWBM 74728
USNM B5659
MNHN 2-22
ZMUC 119500
ANWC 26897
MNHN CG 1998-823

Australia
Mauritius
Solomon Islands
Australia
Australia
Australia
Tanimbar
Australia
MakiraSulawesi
British Samoa
Mindanao
Philippines
British Samoa
Roti, SW of Timor
Australia
Congo
Australia
Kenya
Australia
Italy
New Guinea
Australia
Australia
New Guinea
RussiaMyanmar
Cameroon
Kenya
New Guinea
Malawi

HM002920
HM002921
EF052799
EU380474
EF441216
DQ406669
HM002922
EF052814
HM002923
EU380477

HM003019
HM003020
EU380433

HM002987
HM002988
EF052774

EF441238
EU272119
HM003021
EU380438
HM003022
EU380439

HM002945
HM002946
EF052826
EU380505
AY064732
AY064741
HM002947
EF052840
HM002948
EU380507

EU273381
EF052815

EU273361
EU380440

EU273403
EF052841

HM002924
EF052816
HM002925

HM003023
EU380441
HM003024

MVF 722

Australia

EF052755
EU273389
EF441222
EU273385

EU273363
EU273370
EF441244
EU273366

EF052818
EF052754
DQ406658
DQ406654

EU380450
EU380451

DQ406648

EU380413

HM002949
EF052842
HM002950
FJ821090
AY529926
FJ821094
EF052766
EU273411
AY064728
EU273407
FJ821099
EF052843
EF052765
AY529930
AY529931
EU273416
AY529912

EF441220

EF441242

AY064744

DQ406651

EU380457

AY064749
AY064756
HM002989
EF052785
HM002990
HM002991
HM002992
HM002993
EF052786
HM002994
HM002995
EF052787
HM002996
AY529961
EF052693
GQ494087
AY064755
EU600813
EF052788
EF052692
AY529965
AY529966
AY529947
AY064754

AMNH, American Museum of Natural History, USA; ANSP, Academy of Natural Science, Philadelphia, USA; ANWC, Australian National Wildlife Collection, Canberra, Australia; BMNH, British Museum of Natural History, Tring, England; FMNH, Field Museum of Natural History, Chicago,
USA; MCSNC, Museo Civico di Storia Naturale di Carmagnola, Italy; MNHN, Muséum National d¢Histoire Naturelle, Paris, France; MV, Museum
Victoria, Australia; NRM, Naturhistoriska Riksmuseet, Stockholm, Sweden; RB, Rauri Bowie, Museum of Vertebrate Zoology (MVZ), University ofCalifornia, Berkeley; RG, Musée Royal de l’Afrique Centrale, Tervuren, Belgium; RMNH, Rijksmuseum van Natuurlijke Histoire, Leiden, The
Netherlands; SAM = South Australian Museum, Australia; SEOR, Société d’Etudes Ornithologiques de la Réunion; USNM, United States National
Museum (Smithsonian), Washington DC, USA; UWBM, University of Washington, Burke Museum, Seattle, USA; ZMUC, Zoological Museum,
University of Copenhagen, Denmark. All ingroup taxa are vouchered.
GAPDH, glyceraldehyde-3-phosphodehydrogenase; ODC, ornithine decarboxylase; Myo2, myoglobin intron-2; ND2, NADH dehydrogenase subunit 2.

distribution patterns (Ronquist, 1997), thus it has proven to bean appropriate and popular biogeographical reconstruction
method. A limitation of the implementation of diva (andmany other biogeographical optimization methods) is that it
requires fully bifurcated trees, and that the method does nottake into account the uncertainty in phylogenetic inference.
In contrast, ‘Bayes-diva’ takes topological uncertainty into
account. Rather than using a single fully resolved topology,
which often is possible only by making certain assumptions
about relationships in a given tree, we sampled 15,000 trees (bythinning the chain, i.e. sampling every nth generation) from
the MCMC output and ran diva on all of them. The frequency
Journal of Biogeography 37, 1767–1781ª 2010 Blackwell Publishing Ltdof ancestral areas for clades was then recorded and plotted as
marginal distributions on the majority-rule consensus tree
derived from the MCMC. The major advantage of the Bayesdiva method is that the marginal distributions for thealternative ancestral areas at each node in the tree are theproduct of the phylogenetic uncertainty in the rest of the tree
and the uncertainty in the biogeographical reconstruction of
the node of interest.
In this study, which focuses on members of the coreCampephagidae, we included several other groups known to be
the closest relatives to Campephagidae, such that the basal partof core Campephagidae is no longer the root of the tree. This is1771

K. A. Jønsson et al.
because ancestral reconstructions become increasingly unreliable towards the root of the tree, which can cause the ancestral
distributions at the root to include all areas analysed (Ronquist, 1996). Ancestral reconstructions depend on both thenodes above (more distally) and below (more basally) thegroup of interest.
We assigned seven geographical areas for the diva analysis
considering evidence of historical relationships of geologicalplates and terranes (Audley-Charles, 1981; Hall, 1998; Moss &Wilson, 1998): A) Australia/New Guinea (including the
Bismarcks and the Admirality Islands, which, given their
immediate proximity to New Guinea, have been easilycolonizable in recent time); B) Wallacea (the area east of
Borneo and Bali and west of New Guinea); C) the Pacificislands; D) the Philippine islands; E) Asia (including Sumatra,
Borneo and Java, which are part of the Asian plate and havebeen connected to the Asian mainland on multiple occasions);
F) Africa (including Madagascar); and G) the Indian Ocean
islands of Réunion and Mauritius. The analysis was carried out
constraining the maximum number of areas encompassed by
the ancestral distributions to the maximum size of extant
ranges using the maxareas (=2) option in diva. This isequivalent to assuming that the ancestors of the group inquestion have the same ability to disperse as their extant
descendants and therefore that ancestral ranges were similarin size to extant ranges (Sanmartı́n, 2003; Nylander et al.,2008b).
Dating analyses
We used beast v.1.4.6 (Drummond et al., 2002, 2006;
Drummond & Rambaut, 2007) to estimate the divergencedates within the Campephagidae. We assigned the best fitting
model, as estimated by MrModeltest2, to each of thepartitions. Because there are no fossils within or close to theCampephagidae, we used a combination of the island age ofRéunion (2 Ma; see below) and a rate of 0.028 substitutions
per site per lineage per million years for ND2 (corrected
pairwise distances), which is derived from Galapagos mockingbirds (Drovetski et al., 2004) and secondary calibration
points from Barker et al. (2004). We assumed a YuleSpeciation Process for the tree prior and an uncorrelated
lognormal distribution for the molecular clock model (Drummond et al., 2006; Ho, 2007). We used default prior distributions for all other parameters and ran MCMC chains for 50
million generations. The analysis was repeated three times. We
used the program Tracer (Rambaut & Drummond, 2007) toassess convergence diagnostics.
Calibration points
The use of geological calibration points has been applied
successfully for several avian groups, such as scops-owls (Fuchs
et al., 2008), sunbirds (Warren et al., 2003) and whistlers(Jønsson et al., 2010a). A general problem when using islands
as calibration points is that colonization may have taken place
1772

long after their emergence. Thus, in order to use the age of anisland as a calibration point, the island should be fairly young
and as remote as possible. In the present study, the only islands
that fit these criteria are Mauritius (7.8 Myr old; McDougall &Chamalaun, 1969) and Réunion (c. 2 Myr old; Chevallier &Vatin-Perignon, 1982) in the Indian Ocean. It is unknown
what the error margins for these age estimates are, and using
these exact ages would indeed be misleading. Therefore weapplied to our data a recently published rate extrapolation(2.8% Myr)1) of evolution in ND2 for another family of
passerine birds (Norman et al., 2007). This provides an age
estimate for the split between Coracina newtoni (Réunion) andCoracina typica (Mauritius) at 1.35 Ma. Given the younger ageof Réunion compared with Mauritius, it makes sense to
assume that Mauritius was colonized before the emergence ofRéunion, and that Mauritius would have acted as the sourceof colonists to Réunion once it emerged. So the combinationof island age and ND2 evolution suggests that the split ofthese two species of cuckoo-shrike took place sometimearound 1.5 Ma. Thus, for this calibration we chose a normally
distributed prior at 1.5 Ma ± 0.25 SD (age within 95%
confidence intervals (CI) = 1.089–1.911 Ma). Because it isimportant to have calibration points both basally and distallyin the phylogenetic tree, we also used secondary calibration
points from Barker et al. (2004). Barker et al. (2004) used a
nonparametric rate-smoothing (NPRS) and a penalized-likelihood (PL) approach, as well as a calculation based on the
DNA–DNA hybridization study of Sibley & Ahlquist (1990).
The split of Acanthisitta from all other passerines is estimatedat c. 82 Ma and was used to calibrate the tree. We used the ageof two of the oscine splits (nodes 7 and 11 in Barker et al.,2004). We used an average of the different age estimatespresented by Barker et al. (2004). This translates into an
estimate of the age of the split between Artamus andMalaconotus to be 27.5 Ma ± 1 SD (age within 95%
CI = 25.86–29.14 Ma) and the split between Menura and allother oscines at 62.5 Ma ± 1 SD (age within 95% CI = 60.86–
64.46 Ma).
Our chronogram can also be visualized as a relative-timechronogram (when removing the absolute age estimates). Thisis important because we are also interested in assessing
whether dispersal events of certain clades are centred on aparticular time that could be explained by discrete palaeogeographical events.
RESULTS
Phylogenetic analyses
The use of old museum specimens at times made amplificationof some DNA segments rather difficult, thus a few sequence
fragments are missing (Table 1).
For GAPDH we sequenced between 272 and 298 bp, but
managed to obtain only 215 bp from Aleadryas rufinucha,
197 bp from Coracina analis, 197 bp from Coracina bicolor,
197 bp from Coracina caledonica, 197 bp from Coracina
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Phylogeny and biogeography of cuckoo-shrikes
graueri, 197 bp from Coracina leucopygia, 235 bp from Coracina longicauda, 197 bp from Coracina melanoptera, 180 bp
from Coracina montana, 197 from Coracina ostenta, 163 bp
from Coracina tenuirostris tenuirostris, 197 bp from Lalage
sueurii, and 197 bp from Lobotos oriolinus. The total GAPDHalignment was 321 bp.
For ODC intron-6 and intron-7 we obtained 540–615 bp,
but managed to obtain ODC intron-6 (245–249 bp) only
from Coracina analis, Coracina bicolor, Coracina leucopygia,
Coracina ostenta, Lalage leucopygialis and Lalage sueurii. The
total alignment of ODC intron-6 and intron-7 was 634 bp.
For myoglobin intron-2 we sequenced 669–708 bp; however, some of the sequences (mostly some accessed from
GenBank) were shorter, ranging between 625 and 662 bp.
These include Telophorus sulfureopectus, Prionops retzii, Platysteira cyanea, Malaconotus blanchoti, Artamus cyanopterus,
Coracina novaehollandiae, Coracina pectoralis, Lalage tricolorand Batis poensis. The total alignment of myoglobin intron-2
was 716 bp.
For NADH dehydrogenase subunit 2 we typically obtained
between 1023 and 1041 bp, but only 1013 bp from Coracina pectoralis, 960 bp from Coracina boyeri, 919 bp from
Coracina polioptera, 827 bp from Coracina typica, 816 bp from
Coracina newtoni, 772 bp from Campochaera sloetti, Coracina
bicolor, Coracina striata kochii and Coracina temminckii,
669 bp from Coracina ceramensis, Coracina ostenta andCoracina sula, and 525 bp from Coracina dispar.
Analyses performed on the concatenated data set (sixpartitions: GAPDH, ODC, Myo2, 1st, 2nd, 3rd mtDNA codon
positions; maximum likelihood (ML): )ln 30273.75, Bayesianinference (BI) harmonic mean )ln 28990.66) and on theindividual partitions (GAPDH: AIC: HKY + C, ML: )ln
2177.63, BI harmonic mean )ln 2447.96; ODC: AIC: GTR + C,
ML: )ln 3231.03, BI harmonic mean )ln 3567.96; Myo2: AIC:HKY + C, ML: )ln 3767.67, BI harmonic mean )ln 4163.17;
ND2: AIC: TVM + C, ML: )ln 19497.81, BI harmonic mean)ln 19178.53) yielded 50% majority-rule consensus treesthat were topologically congruent for well-supported nodes(posterior probability >0.95 and bootstrap values >70%). The
nuclear gene trees (GAPDH, ODC and Myo2) (see Appendices S1–S3 in Supporting Information) provide only a few
well-supported clades. This was not unexpected, and reflects
the fact that the nuclear genes used evolve too slowly to resolve
closely related young species within the Campephagidae. The
nuclear data do, however, provide evidence for the partitioningof some basal clades. The ND2 gene tree (Appendix S4)
provides better resolution in the distal part of the tree, and the
combined analysis (Fig. 1) of both mitochondrial and nuclear
genes generates a rather robust and densely sampled coreCampephagidae phylogeny.
Scores of the best likelihood trees were within 0.5 likelihood
units of the best tree recovered in each of the other four garliruns, suggesting that the five runs had converged. The ML tree
topology was completely congruent with the BI topology forwell-supported nodes (posterior probability >0.95 and bootstrap values >70).
Journal of Biogeography 37, 1767–1781ª 2010 Blackwell Publishing Ltd

Bayesian dispersal–vicariance analysis
The Bayes-diva analysis (Fig. 2) recovers the origin of oscines
to be Australian (nodes 1–6) in concordance with several
other studies (Christidis, 1991; Barker et al., 2002; Edwards &Boles, 2002; Ericson et al., 2002). The origin of the Campephagidae (node 7) including Pericrocotus, which occurs inAsia (including Borneo, Sumatra and Java), is also recovered
as Australian/Asian (AE; >0.97 CI). Given that the origin of
the next node (8) is Australian (A; >0.96 CI) and that nodes(9 and 10) further up in the tree are Australian/African inorigin (AF; >0.99 and >0.96 CI, respectively), it is mostparsimonious to assume that the initial dispersal event from
Australia led to Pericrocotus colonizing Asia, where the genus
subsequently radiated. Pericrocotus is distributed in mainland
Asia, Borneo, Java, Sumatra and Palawan, but is absent from
Wallacea (Jønsson et al., 2010b). The absence of Pericrocotus
from Wallacea suggests that direct dispersal from Australia toAsia, not island-hopping across the Wallacean archipelago, isthe most likely scenario. The core Campephagidae, however,remained in Australo-Papua. Node 10 splits into an allAfrican group (four species) and another group more or lessrestricted to Australo-Papua with a few rather late dispersals
to Asia. Node 9 also splits into an all-African group (three
species) and a more heterogeneous group consisting of species
belonging basally to Australo-Papua, Wallacea, the Philippines
and the Pacific. It is noteworthy that there is no indication of
lineage sharing between Australo-Papua and mainland Asia at
this stage. Furthermore, there is a rather clear pattern ofdispersal between Australia and Africa, bypassing Asia. Apartfrom Pericrocotus, there is no dispersal to Asia from AustraloPapua in the basal part of the Campephagidae phylogeny. In
fact, all Asian members of the core Campephagidae are foundat terminal points in the phylogeny. Ancestral area reconstructions in distal parts of the tree become somewhat
ambiguous, probably due to a high exchange of species
between various regions leading to an obscured biogeographical signal. It should be noted, though, that the two Indian
Ocean species (Coracina newtoni and Coracina typica) have an
Asian rather than an African origin.
Dating analyses
The results of the beast dating analysis (Fig. 3) indicate theorigin of the Campephagidae to be in the Late Oligocene and theorigin of the core Campephagidae to be in the Early Miocene. We
are particularly interested in the timing of dispersal between
Australia and Africa. Thus, the time of common ancestry for the
African clade comprising Coracina cinerea, C. graueri, C.
pectoralis and C. caesia, and its sister-group is found to be
14 Ma (95% highest posterior density (HPD) = 11.5–17 Ma)
and the time of common ancestry for the African clade
comprising Campephaga and Lobotos and its sister-group is
17 Ma (95% HPD = 14–20 Ma). Coracina azurea is a species
that proves difficult to place in the phylogeny. Although C.
azurea is sister to C. abbotti in both the Bayesian and the beast
1773

K. A. Jønsson et al.

Figure 1 The 50% majority rule consensus tree of the core Campephagidae and close relatives obtained from the Bayesian analysis of the
combined data set: glyceraldehyde-3-phosphodehydrogenase (GAPDH), ornithine decarboxylase (ODC), myoglobin intron-2 (Myo2),
NADH dehydrogenase subunit 2 (ND2). Support values are indicated to the left of the nodes. Above the branch is the posterior probability(only values above 0.90 are shown, asterisks indicate posterior probabilities of 1.00). Below (or above to the right of) the branch is the
maximum likelihood bootstrap value (only values above 70% are shown) from 500 pseudoreplicates. Terminal taxa distributed in Africa,
light grey; terminal taxa distributed in Asia (and the Indian Ocean), dark grey. All other taxa are either Australo-Papuan, Wallacean,Philippine or Pacific in distribution.
Figure 2 A summary of the Bayesian dispersal–vicariance analysis (Bayes-diva) of the core Campephagidae and close relatives. The tree is achronogram based on a 50% majority-rule consensus tree of a Bayesian Markov chain Monte Carlo (MCMC) analysis of a combined data setof mitochondrial [NADH dehydrogenase subunit 2, (ND2)] and nuclear [glyceraldehyde-3-phosphodehydrogenase (GAPDH), ornithine
decarboxylase (ODC) and myoglobin intron-2 (Myo)] DNA sequences. Breeding region for each taxon, as delimited in the map, isgiven after each taxon name. Pie charts at internal nodes represent the marginal probabilities for each alternative ancestral area derived using
diva while integrating over tree topologies using MCMC. These probabilities account for the phylogenetic uncertainty in the rest of thetree and the biogeographical uncertainty (multiple equally parsimonious reconstructions) at each node, conditional on this node occurringin a given sampled topology. In the pie charts, the first four areas with highest probability are coloured according to relative probability inthe following order: white > red > blue > grey, and any remaining areas are collectively coded in black. Some basal nodes have been
assigned a number, which is referred to in the Discussion. Biogeographical regions: A, Australia/New Guinea; B, Wallacea; C, The PacificArchipelago; D, The Philippine Archipelago; E, Asia (including Sumatra, Java and Borneo); F, Africa; G, Mauritius/Réunion.

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Phylogeny and biogeography of cuckoo-shrikes

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1775

K. A. Jønsson et al.
Oreoica gutturalis
Aleadryas rufinucha
Ornorectes cristatus
Pericrocotus divaricatus
Pericrocotus erythropygius
Campochaera sloetti
Lalage leucomela
Lalage atrovirens
Lalage melanoleuca
Lalage leucopygialis
Lalage nigra
Coracina melanoptera
Coracina fimbriata
Coracina polioptera
Coracina melaschista
Coracina newtoni
Coracina typica
Lalage maculosa
Lalage leucopyga
Lalage sueurii
Lalage tricolor
Lalage sharpei
Coracina montana
Coracina dispar
Coracina dohertyi
Coracina ceramensis
Coracina tenuirostris admiralitatis
Coracina salomonis
Coracina morio
Coracina incerta
Coracina tenuirostris matthiae
Coracina tenuirostris heinrothi
Coracina remota
Coracina sula
Coracina t. tenuirostris
Coracina holopolia
Coracina mindanensis
Coracina ostenta
Coracina coerulescens
Coracina analis
Coracina az urea
Coracina abbotti
Coracina mcgregori
Lobotos oriolinus
Campephaga petiti
Campephaga flava
Coracina cinerea
Coracina graueri
Coracina pectoralis
Coracina caesia
Coracina caeruleogrisea
Coracina temminckii
Coracina max ima
Coracina bicolor
Coracina l. lineata
Coracina lineata sublineata
Coracina boyeri
Coracina welchmani
Coracina papuensis artamoides
Coracina atriceps
Coracina papuensis sclaterii
Coracina ingens
Coracina striata guillemardi
Coracina macei
Coracina striata kochii
Coracina leucopygia
Coracina scistacea
Coracina larvata
Coracina caledonica
Coracina novaehollandiae
Coracina longicauda
Peltops blainvillii
Gymnorhina tibicen
Artamus cyanopterus
Machaerirhynchus flaviventer
Malaconotus blanchoti
T elophorus sulfureopectus
Batis poensis
Platysteira cyanea
Prionops retz ii
Rhagologus leucostigma
Pachycephala pectoralis
Daphoenositta chrysoptera
Oriolus oriolus
Eopsaltria australis
Orthonyx temminckii
Menura novaehollandiae

Paleocene
60

Oligocene

Eocene
50

40

30

20

Miocene10

PP

0

Figure 3 Chronogram of the core Campephagidae and close relatives based on the beast analysis. Blue bars represent 95% highest
posterior density (HPD) intervals. Terminal taxa are coloured according to present distributions: yellow, Australia/New Guinea (including
the Bismarck Archipelago); green, Wallacea; red, Africa; blue, Pacific Archipelago; purple, Philippine Archipelago; brown, Asia (including
Sumatra, Java, Borneo); black, Mauritius and Réunion. The vertical red column indicates the time span when the three African clades
dispersed from Australia to Africa; shaded red indicates 95% HPD intervals; red dotted line indicates putative dispersal between Australia
and Africa in the Oligocene.

analyses, the relationship has no support and thus should beconsidered unresolved. Therefore, the split between C. azurea
and any sister taxon is cautiously set to 13.5 Ma (95%
HPD = 11–16 Ma). With these age estimates, it seems that thethree dispersal events to Africa have taken place in a rather
narrow time frame in the Miocene between 13.5 and 17 Ma(95% HPD = 11–20 Ma). Dispersal into mainland Asia, on theother hand, did not take place before the Pliocene.
DISCUSSION
Systematics and taxonomy
Our study presents the first robust phylogeny of the coreCampephagidae (Campephagidae with the exclusion of
1776

Pericrocotus) including all genera, and lacks only a few species
with restricted ranges (mainly on smaller islands in thearchipelagos surrounding Australo-Papua). The missing island
taxa are generally linked to taxa on larger nearby landmasses(Taylor, 2005). Therefore, we feel confident in recommending
some preliminary taxonomic changes. Our study confirms themonophyly of the Campephagidae, with a core Campephagidae group consisting of the genera Coracina, Lalage, Campephaga, Lobotos and Campochaera. These genera, however, are
not monophyletic. Two primary clades are apparent, as also
noted by Fuchs et al. (2007) and Jønsson et al. (2008). One
clade (A) comprises a subset of Coracina Vieillot, 1816
(including the type species for the genus, Coracina papuensis)
and thus should maintain this name. Clade B contains speciesof two genera, Lalage and Coracina. The oldest name for this
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Phylogeny and biogeography of cuckoo-shrikes
assemblage is Lalage (Lalage Boie, 1826), and thus we proposethat the whole clade be treated under the name Lalage.
Whether or not the genus Lobotos, which is sister to the genus
Campephaga, should be maintained as a separate genus, isdebatable. Peters (1960) argued that there is little reason tomaintain the genus Lobotos as it does not seem to differ
substantially in morphology from Campephaga. However,mitochondrial ND2 sequence data differ by 11.62% between
the genera, perhaps suggesting that genus status is valid. Basedon the results of the present study, the family Campephagidae
now contains five genera: Pericrocotus, Coracina, Campephaga(including Lobotos), Campochaera and Lalage (including
several species previously assigned to Coracina) (Fig. 1).
Biogeography
The Bayes-diva analysis indicates that the origin of the basal
clades of oscine passerine birds (represented here by Menura
and Orthonyx nodes 1 and 2), Petroicidae (represented by
Eopsaltria node 3), and the basal part of the core Corvoidea(represented by all other taxa outside Campephagidae nodes
4–6) is Australo-Papuan. The Campephagidae (node 7),
including Pericrocotus, has an Australian/Asian origin, whereas
the origin of the core Campephagidae (node 8) is AustraloPapuan. This could be interpreted in one of two ways: (1)members of the Campephagidae were historically distributed
throughout the Australo-Papuan and Asian region and later
retracted to Australo-Papua when the core Campephagidae
started radiating, or (2) a Pericrocotus ancestor dispersed out ofAustralia and colonized Asia as the first dispersal event within
this group. We argue that the latter is the most likely scenario,
given that there are no other Asian species in the basal part ofthe core Campephagidae phylogeny. At the time when
Pericrocotus diverged from the core Campephagidae some
25 Ma, there was still a substantial body of water between
Australia and Asia, and it is more parsimonious to assume thata single proto-Pericrocotus colonized Asia (which is also
supported by the Bayes-diva analysis) than to assume ahistorical range expansion followed by a range retraction of thewhole Campephagidae. A molecular study of Pericrocotus
unequivocally demonstrates that the radiation is centred inmainland Asia, from where dispersal took place in a southeastwards direction to the Indonesian archipelago (Jønsson
et al., 2010b). This dispersal pattern supports the idea that
Pericrocotus colonized Asia by long-distance dispersal rather
than by island hopping from Australia across Wallacea.
African species occur in two distinct basal clades within thecore Campephagidae. One clade contains Campephaga and
Lobotos. Another clade contains four species: Coracina caesia,
C. pectoralis, C. graueri and C. cinerea. Coracina azurea isdifficult to place, but is linked with low support to C. abbotti of
Sulawesi. As C. azurea is not closely related to other Africantaxa, it seems to constitute an independent colonization of
Africa at a relatively early time in the evolution of the coreCampephagidae. Thus, we find three colonizations of Africa,
none of which appears to be closely linked to the two very
Journal of Biogeography 37, 1767–1781ª 2010 Blackwell Publishing Ltd

distal clades of mainland Asian taxa. This indicates that theCampephagidae did not colonize Africa via Asia, unless one
assumes extinction of Asian representatives from all threedispersing clades.
To sum up, we find, based on the phylogeny and the Bayesdiva analysis, that the Campephagidae arose in AustraloPapua, from where one early dispersal event of Pericrocotus
colonized mainland Asia. Africa was colonized twice (perhaps
three times) directly from Australia at around 14–20 Ma.
Within the past 10 Myr, no cuckoo-shrike has colonized
Africa. Rather, within this time frame, there have been anumber of multi-directional dispersal events taking place from
Australo-Papua to the Pacific Islands, the Philippines, Wallacea and mainland Asia. The Indian Ocean Islands of Réunionand Mauritius were among the most recently colonized islands,
and were colonized from Asia rather than Africa during the
Plio/Pleistocene.
Given that there is no evidence for the core Campephagidae
having colonized Asia at an early point, we find little reason tosuspect that cuckoo-shrikes colonized Africa via Asia, as thiswould require numerous extinction events. On the other hand,at the time of the ‘African colonization’ in the mid-Miocene,
Australia had collided with Asia, providing a stepping-stonecolonization pathway across Wallacea; and from the Oligocene
to the mid-Miocene, large expanses of forest covered acontinuous area from Southeast Asia to Africa, creating adispersal corridor for forest-associated taxa (Janis, 1993). Partsof this region became significantly drier towards the end of the
Miocene and caused parts of western Asia, the Arabian
Peninsula and north-eastern Africa to become arid grasslandand desert (Jacobs et al., 1999). With these climatic andtectonic changes in mind, it seems by no means impossiblethat members of the core Campephagidae dispersed via islands
and forested land masses from Australia to Africa, and that
Asian members subsequently went extinct. However, it does
seem difficult to explain that Pericrocotus successfully radiatedand still has many extant members throughout Asia, whereas
all members of the core Campephagidae that expanded in the
Miocene went extinct.
A number of recent studies have demonstrated transoceanic
dispersal as a possible mechanism to explain present disjunctdistribution patterns of various groups of vertebrates (Yoder
et al., 2003; Heinicke et al., 2007; Pereira et al., 2007).
However, the evolutionary link between Australo-Papua andAfrica/Madagascar remains rather unusual, although it has
been suggested for several angiosperm families (Baum et al.,
1998; Davis et al., 2002; Mummenhoff et al., 2004). We knowof only few terrestrial avian examples for which a link between
Australia and Africa has been suggested (Fuchs et al., 2006;
Jønsson & Fjeldså, 2006; Jønsson et al., 2008; Wright et al.,2008; Schweizer et al., 2010). For the Psittaciformes (Wrightet al., 2008; Schweizer et al., 2010) the authors reach somewhat
different conclusions regarding dispersal. It seems that Psittaciformes is rather old, and that the link could be explained byvicariance events associated with the break-up of Gondwana
and perhaps some dispersal events over relatively short1777

K. A. Jønsson et al.
distances. Thus, the present study is the most substantiatedavian example of long-distance ocean dispersal and colonization of Africa across the Indian Ocean from Australo-Papua.
Jønsson & Fjeldså (2006) have previously suggested thatland plateaux in the southern Indian Ocean could havefacilitated dispersal from Australia to Africa at around 45–
50 Ma. However, by the Miocene these large land plateaux had
subsided markedly (Coffin et al., 2002; Wallace et al., 2002).
Although the Kerguelen Archipelago had several subaerial
islands, it was far to the south and is unlikely to have aided
birds crossing the Indian Ocean. Rather, it seems that cuckooshrikes dispersed directly across the Indian Ocean to Africa
during two or three individual colonizations within a narrow
time frame between 13.5 and 17 Ma.
The majority of cuckoo-shrike species are rather sedentary
forest-associated species, and the few real migratory members
inhabit high-latitude regions in continental Asia. However, thegeneral lack of movements probably reflects the fact that thebirds inhabit regions where environmental conditions vary
little by season. Clearly, cuckoo-shrikes disperse well because
they have colonized isolated oceanic islands in Micronesia,
Melanesia and the Indian Ocean (Taylor, 2005). It seems thatmobility is very plastic among birds, probably best exemplified
by another avian family, Rallidae, whose members are generally
thought of as poor flyers but nevertheless have colonized amultitude of remote islands and archipelagos across the globe(Taylor, 1996).
Although other avian studies in the Indian Ocean havedemonstrated that transoceanic dispersal is important forcolonizing islands (Warren et al., 2003; Fuchs et al., 2008),
the present study infers the occurrence of transoceanic
dispersal over an unusually large body of water (the Indian
Ocean). A recent study of Turdus thrushes found evidence
for direct dispersal from Africa to the Caribbean (Voelker
et al., 2009), lending further support to the idea that longdistance ocean dispersal may be an important biogeographical force.
ACKNOWLEDGEMENTS
We are grateful to the following institutions for granting accessto toe-pad, blood and tissue samples: AMNH, American
Museum of Natural History, USA; ANSP, Academy of Natural
Science, Philadelphia, USA; ANWC, Australian National
Wildlife Collection, Canberra, Australia; BMNH, British
Museum of Natural History, Tring, England; FMNH, Field
Museum of Natural History, Chicago, USA; MCSNC, Museo
Civico di Storia Naturale di Carmagnola, Italy; MNHN,
Muséum National d¢Histoire Naturelle, Paris, France; MV,
Museum Victoria, Australia; NRM, Naturhistoriska Riksmuseet, Stockholm, Sweden; RG, Musée Royal de l’AfriqueCentrale, Tervuren, Belgium; RMNH, Rijksmuseum van
Natuurlijke Histoire, Leiden, The Netherlands; SAM = South
Australian Museum, Australia; SEOR, Société d’Etudes Ornithologiques de la Réunion; USNM, United States National
Museum (Smithsonian), Washington DC, USA; UWBM,
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University of Washington, Burke Museum, Seattle, USA; andZMUC, Zoological Museum, University of Copenhagen,Denmark. We would like to thank Walter Boles for taxonomic
advice. K.A.J. would also like to thank Manickchand Puttoo,
Director and Senior Research and Development Officer andRajen Sookhareea of the National Parks and Conservation
Service for kindly issuing a collecting permit in Mauritius. In
the field K.A.J. was assisted by Lone Raffray of the Mauritian
Wildlife Foundation and volunteers Anna Reuleaux, Jannie
Fries Linnebjerg, Elaine Fraser, Jason van de Wetering andrangers Paul and Mario, who all proved indispensable. K.A.J.
also thanks Thomas Ghestemme for his help in the field. K.A.J.
would also like to acknowledge support from the AustralianMuseum Postgraduate Awards 2006/07.
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Journal of Biogeography 37, 1767–1781ª 2010 Blackwell Publishing Ltd

SUPPORTING INFORMATION
Additional supporting information may be found in the online
version of this article:
Appendix S1 The 50% majority rule consensus tree of thecore Campephagidae obtained from the Bayesian analysis ofglyceraldehyde-3-phosphodehydrogenase (GAPDH).
Appendix S2 The 50% majority rule consensus tree of thecore Campephagidae obtained from the Bayesian analysis ofornithine decarboxylase (ODC).
Appendix S3 The 50% majority rule consensus tree of thecore Campephagidae obtained from the Bayesian analysis of
myoglobin intron-2 (Myo2).
Appendix S4 The 50% majority rule consensus tree of thecore Campephagidae obtained from the Bayesian analysis of
NADH dehydrogenase subunit 2 (ND2).
As a service to our authors and readers, this journal provides
supporting information supplied by the authors. Such materials are peer-reviewed and may be re-organized for online
delivery, but are not copy-edited or typeset. Technical support
issues arising from supporting information (other than
missing files) should be addressed to the authors.

BIOSKETCHES
Knud A. Jønsson is a PhD student at the ZoologicalMuseum, University of Copenhagen, Denmark. His maininterests include the systematics and historical biogeography ofpasserine birds with a particular emphasis on the Indo-Pacificregion.
Jon Fjeldså is a professor at the Zoological Museum,
University of Copenhagen, Denmark. He has broad interestsin the phylogeny and systematics of birds as well as a generalinterest in several other aspects of ornithology, ecology andconservation.
Author contributions: K.A.J. and J.F. conceived the ideas;
K.A.J. and J.A.A.N. collected and analysed the data. All authors
took part in the writing process.

Editor: Lawrence Heaney

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