A SINGLE ANCIENT ORIGIN OF BROOD PARASITISM IN AFRICAN FINCHES: IMPLICATIONS FOR HOST‐PARASITE COEVOLUTION (original) (raw)

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Department of Biology, Boston University, Boston, Massachusetts 02215

Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48109

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Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48109

Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, Michigan 48109

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Published:

01 December 2001

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Abstract

Abstract Robust phylogenies for brood‐parasitic birds, their hosts, and nearest nesting relatives provide the framework to address historical questions about host‐parasite coevolution and the origins of parasitic behavior. We tested phylogenetic hypotheses for the two genera of African brood‐parasitic finches, Anomalospiza and Vidua, using mitochondrial DNA sequence data from 43 passeriform species. Our analyses strongly support a sister relationship between Vidua and Anomalospiza, leading to the conclusion that obligate brood parasitism evolved only once in African finches rather than twice, as has been the conventional view. In addition, the parasitic finches (Viduidae) are not recently derived from either weavers (Ploceidae) or grassfinches (Estrildidae), but represent a third distinct lineage. Among these three groups, the parasitic finches and estrildids, which includes the hosts of all 19 Vidua species, are sister taxa in all analyses of our full dataset. Many characters shared by Vidua and estrildids, including elaborate mouth markings in nestlings, unusual begging behavior, and immaculate white eggs, can therefore be attributed to common ancestry rather than convergent evolution. The host‐specificity of mouth mimicry in Vidua species, however, is clearly the product of subsequent host‐parasite coevolution. The lineage leading to Anomalospiza switched to parasitizing more distantly related Old World warblers (Sylviidae) and subsequently lost these characteristics. Substantial sequence divergence between Vidua and Anomalospiza indicates that the origin of parasitic behavior in this clade is ancient (∼20 million years ago), a striking contrast to the recent radiation of extant Vidua. We suggest that the parasitic finch lineage has experienced repeated cycles of host colonization, speciation, and extinction through their long history as brood parasites and that extant Vidua species represent only the latest iterations of this process. This dynamic process may account for a significantly faster rate of DNA sequence evolution in parasitic finches as compared to estrildids and other passerines. Our study reduces by one the tally of avian lineages in which obligate brood parasitism has evolved and suggests an origin of parasitism that involved relatively closely related species likely to accept and provide appropriate care to parasitic young. Given the ancient origin of parasitism in African finches, ancestral estrildids must have been parasitized well before the diversification of extant Vidua, suggesting a long history of coevolution between these lineages preceding more recent interactions between specific hosts and parasites.

Literature Cited

Aragon

,

S.

,

A. P.

Møller

,

J. J.

Soler

, and

M.

Soler

.

1999

.

Molecular phylogeny of cuckoos supports a polyphyletic origin of brood parasitism

.

J. Evol. Biol.

12

:

495

–

506

.

Avise

,

J. C.

,

D.

Walker

, and

G. C.

Johns

.

1998

.

Speciation durations and Pleistocene effects on vertebrate phylogeography

.

Proc. R. Soc. Lond. B

265

:

1707

–

1712

.

Bannerman

,

D. A.

1949

. The birds of tropical West Africa. Vol.

7

.

Crown Agents

, London.

Baptista

,

L. F.

1973

.

Song mimesis by a captive Gouldian finch

.

Auk

90

:

891

–

894

.

Bentz

,

G. D.

1979

.

The appendicular myology and phylogenetic relationships of the Ploceidae and Estrildidae (Aves: Passeriformes)

.

Bull. Carnegie Mus. Nat. Hist.

15

:

1

–

25

.

Birkhead

,

T. R.

,

T.

Burke

,

R.

Zann

,

F. M.

Hunter

, and

A. P.

Krupa

.

1990

.

Extrapair paternity and intraspecific brood parasitism in wild zebra finches, Taeniopygia guttata, revealed by DNA fingerprinting

.

Behav. Ecol. Sociobiol.

27

:

315

–

324

.

Bremer

,

K.

1988

.

The limits of amino‐acid sequence data in angiosperm phylogenetic reconstruction

.

Evolution

42

:

795

–

803

.

Brown

,

W. M.

,

E. M.

Prager

,

A.

Wang

, and

A. C.

Wilson

.

1982

.

Mitochondrial DNA sequences of primates: the tempo and mode of evolution

.

J. Mol. Evol.

18

:

225

–

239

.

Campbell

,

B.

, and

E.

Lack

, eds.

1985

. A dictionary of birds.

T. & A. D. Poyser

, Calton, U.K.

Chapin

,

J. P.

1917

.

The classification of the weaver‐birds

.

Bull. Am. Mus. Nat. Hist.

37

:

243

–

280

.

Chapin

,

J. P.

1954

.

The birds of the Belgian Congo

.

Part 4. Bull. Am. Mus. Nat. Hist. 75B

.

Clayton

,

N. S.

1989

.

Effects of cross‐fostering on selective song learning in estrildid finches

.

Behaviour

109

:

163

–

175

.

Collias

,

N. E.

, and

E. C.

Collias

.

1964

.

Evolution of nest‐building in the weaverbirds (Ploceidae)

.

Univ. Calif. Publ. Zool.

73

:

1

–

162

.

Cooper

,

A.

1994

. DNA from museum specimens. pp.

149

–

165

in

B.

Herrmann

and

S.

Herrmann

, eds. Ancient DNA: recovery and analysis of genetic material from paleontological, archaeological, museum, medical, and forensic specimens.

Springer

, New York.

Crook

,

J. H.

1958

.

Études sur le comportement social de Bubalornis a. albirostris (Vieillot)

.

Alauda

26

:

161

–

195

.

Crook

,

J. H.

1964

.

The evolution of social organization and visual communication in the weaver birds (Ploceinae)

.

Behaviour

10

(

Suppl.

):

1

–

178

.

Davies

,

N. B.

1992

. Dunnock behaviour and social evolution.

Oxford Univ. Press

, Oxford, U.K..

Davies

,

N. B.

2000

. Cuckoos, cowbirds and other cheats.

T. and A. D. Poyser

, London.

Davies

,

N. B.

, and

M. de L.

Brook

.

1988

.

Cuckoos versus reed warblers: adaptations and counteradaptations

.

Anim. Behav.

36

:

262

–

284

.

Delacour

,

J.

, and

F.

Edmond‐Blanc

.

19331934

.

Monographie des veuves (revision des genres Euplectes et Vidua)

.

Oiseau Rev. Fr. Ornithol.

3

:

519

–

562

; 4:52–110.

Desjardins

,

P.

, and

R.

Morais

.

1990

.

Sequence and gene organization of the chicken mitochondrial genome

.

J. Mol. Biol.

212

:

599

–

634

.

Dhindsa

,

M. S.

1983a

.

Intraspecific nest parasitism in two species of Indian weaverbirds Ploceus benghalensis and P. manyar

.

Ibis

125

:

243

–

245

.

Dhindsa

,

M. S.

1983b

.

Intraspecific nest parasitism in the white‐throated munia

.

Notornis

30

:

87

–

92

.

Dhindsa

,

M. S.

1990

.

Intraspecific brood parasitism in the baya weaverbird Ploceus philippinus

.

Bird Behav

8

:

111

–

113

.

Disney

,

H. J. de S.

1980

. Skull pneumatisation in African birds with reference to ageing and breeding and some comparisons with Australian species. pp.

43

–

50

in

D. N.

Johnson

, ed. Proceedings of the fourth pan‐African ornithological congress.

South African Ornithological Society

, Johannesburg, South Africa.

Felsenstein

,

J.

1978

.

Cases in which parsimony and compatibility methods will be positively misleading

.

Syst. Zool.

27

:

401

–

410

.

Felsenstein

,

J.

1985

.

Confidence limits on phylogenies: an approach using the bootstrap

.

Evolution

39

:

783

–

791

.

Fleischer

,

R. C.

,

C. E.

McIntosh

, and

C. L.

Tarr

.

1998

.

Evolution on a volcanic conveyor belt: using phylogeographic reconstructions and K‐Ar based ages of the Hawaiian Islands to estimate molecular evolutionary rates

.

Mol. Ecol.

7

:

533

–

545

.

Foster

,

P. G.

, and

D. A.

Hickey

.

1999

.

Compositional bias may affect both DNA‐based and protein‐based phylogenetic reconstructions

.

J. Mol. Evol.

48

:

284

–

290

.

Freeman

,

S.

1988

.

Egg variability and conspecific nest parasitism in the Ploceus weaverbirds

.

Ostrich

59

:

49

–

53

.

Friedmann

,

H.

1960

. The parasitic weaverbirds. Bulletin no. 223. U.S.

National Museum

, Washington, D.C.

Gibbs

,

H. L.

,

M. D.

Sorenson

,

K.

Marchetti

,

M. de L.

Brooke

,

N. B.

Davies

, and

H.

Nakamura

.

2000

.

Genetic evidence for female host races of the common cuckoo

.

Nature

407

:

183

–

186

.

Giribet

,

G.

, and

W. C.

Wheeler

.

1999

.

On gaps

.

Mol. Phylogenet. Evol.

13

:

132

–

143

.

Gladstein

,

D.

, and

W. C.

Wheeler

.

1996

. POY: phylogeny reconstruction via direct optimization.

American Museum of Natural History

, New York.

Goodwin

,

D.

1960

.

Observations on avadavats and golden‐breasted waxbills

.

Avic. Mag.

66

:

174

–

199

.

Goodwin

,

D.

1971

.

Imprinting, or otherwise, in some cross‐fostered red‐cheeked and blue‐headed cordon‐bleus (Uraeginthus bengalus and U. cyanocephalus)

.

Avic. Mag.

77

:

26

–

31

.

Goodwin

,

D.

1982

. Estrildid finches of the world.

British Museum of Natural History

, London.

Groth

,

J. G.

1998

.

Molecular phylogenetics of finches and sparrows: consequences of character state removal in cytochrome b sequences

.

Mol. Phylogenet. Evol.

10

:

377

–

390

.

Güttinger

,

H. R.

1972

.

Elementwahl und Strophenaufbau in der Gesangsentwicklung einiger Papageiamadinen‐Arten (Gattung: Erythrura, Familie: Estrildidae)

.

Z. Tierpsychol.

31

:

26

–

38

.

Güttinger

,

H. R.

1973

.

Kopiervermögen von Rhythmus und Strophenaufbau in der Gesangsentwicklung einiger _Lonchura_‐Arten (Estrildidae)

.

Z. Tierpsychol.

32

:

374

–

385

.

Heer

,

L.

1996

.

Cooperative breeding by alpine accentors Prunella collaris: polygynandry, territoriality and multiple paternity

.

J. Ornithol.

137

:

35

–

51

.

Hendy

,

M. D.

, and

D.

Penny

.

1989

.

A framework for the quantitative study of evolutionary trees

.

Syst. Zool.

38

:

297

–

309

.

Hudson

,

W. H.

1874

.

Notes on the procreant instincts of the three species of Molothrus found in Buenos Ayres

.

Proc. Zool. Soc. Lond.

(1874):

153

–

174

.

Hughes

,

J. M.

1996

.

Phylogenetic analysis of the cuculidae (Aves, Cuculiformes) using behavioral and ecological characters

.

Auk

113

:

10

–

22

.

Hughes

,

J. M.

2000

.

Monophyly and phylogeny of cuckoos (Aves, Cuculidae) inferred from osteological characters

.

Zool. J. Linn. Soc.

130

:

263

–

307

.

Immelmann

,

K.

1972

.

The influence of early experience upon the development of social behaviour in estrildine finches

.

Proc. Int. Ornithol. Congr.

15

:

316

–

338

.

Immelmann

,

K.

1975

.

Ecological significance of imprinting and early learning

.

Annu. Rev. Ecol. Syst.

6

:

15

–

38

.

Immelmann

,

K.

, and

G.

Immelmann

.

1967

.

Verhaltensökologische Studien an afrikanischen und australischen Estrildiden

.

Zool. Jahrb. Syst. Bd.

94

:

609

–

686

.

Immelmann

,

K.

,

J.

Steinbacher

, and

H. E.

Wolters

.

1965

. Pracht‐finken. Vol.

1

. Astrilde. 2d ed.

Verlag Hans Limberg

, Aachen.

Immelmann

,

K.

,

J.

Steinbacher

, and

H. E.

Wolters

.

1977

. Pracht‐finken. Vol.

2

. Amadinen. 2d ed.

Verlag Hans Limberg

, Aachen.

Jackson

,

W. M.

1993

.

Causes and consequences of conspecific nest parasitism in the northern masked weaver

.

Behav. Ecol. Sociobiol.

32

:

119

–

126

.

Johnsgard

,

P. A.

1997

. The avian brood parasites: deception at the nest.

Oxford Univ. Press

, Oxford, U.K..

Johnson

,

K. P.

, and

S. M.

Lanyon

.

1999

.

Molecular systematics of the grackles and allies, and the effect of additional sequence (cyt b and ND2)

.

Auk

116

:

759

–

768

.

Kilner

,

R. M.

1999

.

Family conflicts and the evolution of nestling mouth colour

.

Behaviour

136

:

779

–

804

.

Kilner

,

R. M.

,

D. G.

Noble

, and

N. B.

Davies

.

1999

.

Signals of need in parent‐offspring communication and their exploitation by the common cuckoo

.

Nature

397

:

667

–

672

.

Klein

,

N. K.

, and

R. B.

Payne

.

1998

.

Evolutionary associations of brood parasitic finches (Vidua) and their host species: analyses of mitochondrial DNA restriction sites

.

Evolution

52

:

566

–

582

.

Klicka

,

J.

, and

R. M.

Zink

.

1997

.

The importance of recent ice ages in speciation: a failed paradigm

.

Science

277

:

1666

–

1669

.

Klicka

,

J.

, and

R. M.

Zink

.

1999

.

Pleistocene effects on North American songbird evolution

.

Proc. R. Soc. Lond. B.

266

:

695

–

700

.

Kroodsma

,

D. E.

, and

J. R.

Baylis

.

1982

. Appendix: a world survey of evidence for vocal learning in birds. pp.

311

–

337

in

D. E.

Kroodsma

and

E. H.

Miller

, eds. Acoustic communication in birds. Vol.

2

.

Academic Press

, New York.

Kumar

,

S.

,

K.

Tamura

, and

M.

Nei

.

1993

.

MEGA: molecular evolutionary genetics analysis

.

Ver. 1.02. Pennsylvania State Univ. University Park, PA

.

Kunkel

,

P.

1969

.

Die Stammesgeschichte der Prachtfinken (Estrildidae) im Lichte des Brutparasitismus der Witwen (Viduinae)

.

Ardea

57

:

172

–

181

.

Lanyon

,

S. M.

1992

.

Interspecific brood parasitism in blackbirds (Icterinae): a phylogenetic perspective

.

Science

255

:

77

–

79

.

Lewis

,

D. M.

1982

.

Cooperative breeding in a population of white‐browed sparrow weavers Plocepasser mahali

.

Ibis

124

:

511

–

522

.

Li

,

W.‐H.

,

T.

Gojobori

, and

M.

Nei

.

1981

.

Pseudogenes as a paradigm of neutral evolution

.

Nature

292

:

237

–

239

.

Lichtenstein

,

G.

2001

.

Selfish begging by screaming cowbirds, a mimetic brood parasite of the bay‐winged cowbird

.

Anim. Behav.

61

:

1151

–

1158

.

Lotem

,

A.

,

H.

Nakamura

, and

A.

Zahavi

.

1992

.

Rejection of cuckoo eggs in relation to host age: a possible evolutionary equilibrium

.

Behav. Ecol.

3

:

128

–

132

.

Lotem

,

A.

,

H.

Nakamura

, and

A.

Zahavi

.

1995

.

Constraints on egg discrimination and cuckoo host coevolution

.

Anim. Behav.

49

:

1185

–

1209

.

Martin

,

A. P.

, and

S. R.

Palumbi

.

1993

.

Body size, metabolic rate, generation time, and the molecular clock

.

Proc. Natl. Acad. Sci.

90

:

4087

–

4091

.

Moran

,

N. A.

1996

.

Accelerated evolution and Muller's rachet in endosymbiotic bacteria

.

Proc. Natl. Acad. Sci.

93

:

2873

–

2878

.

Morel

,

M.‐Y.

1973

.

Contribution à l'étude dynamique de la population de Lagonosticta senegala L

.

(Estrildides) à Richard‐Toll (Sénégal). Interrelations avec le parasite Hypochera chalybeata (Müller) (Viduines). Mémoires du Muséum National d'Histoire Naturelle, Serie A. Zoologie 78, Paris, France

.

Morlion

,

M. L.

1980

.

Pterylosis as a secondary criterion in the taxonomy of the African Ploceidae and Estrildidae

. pp.

27

–

41

in

D. N.

Johnson

, ed.

Proceedings of the fourth pan‐African ornithological congress, South African Ornithological Society, Johannesburg, South Africa

.

Naylor

,

G. J. P.

, and

W. M.

Brown

.

1998

.

Amphioxus mitochondrial DNA, chordate phylogeny, and the limits of inference based on comparisons of sequences

.

Syst. Biol.

47

:

61

–

76

.

Neunzig

,

R.

1929

.

Zum Brutparasitismus der Viduinen

.

J. Ornithol.

77

:

1

–

21

.

Nicolai

,

J.

1964

.

Der Brutparasitismus der Viduinae als ethologisches Problem

.

Z. Tierpsychol.

21

:

129

–

204

.

Nicolai

,

J.

1973

.

Das Lernprogramm in der Gesangsausbildung der Strohwitwe Tetraenura fischeri Reichenow

.

Z. Tierpsychol.

32

:

113

–

138

.

Nunn

,

G. B.

, and

S. E.

Stanley

.

1998

.

Body size effects and rates of cytochrome b evolution in tube‐nosed seabirds

.

Mol. Biol. Evol.

15

:

1360

–

1371

.

Ohta

,

T.

1992

.

The nearly neutral theory of molecular evolution

.

Annu. Rev. Ecol. Syst.

23

:

263

–

286

.

Omland

,

K. E.

,

S. M.

Lanyon

, and

S. J.

Fritz

.

1999

.

A molecular phylogeny of the New World orioles (Icterus): the importance of dense taxon sampling

.

Mol. Phylogenet. Evol.

12

:

224

–

239

.

Pakenham

,

R. H. W.

1939

.

Field notes on the birds of Zanzibar and Pemba

.

Ibis

3

:

522

–

554

.

Payne

,

R. B.

1967

.

Interspecific communication signals in parasitic birds

.

Am. Nat.

101

:

363

–

376

.

Payne

,

R. B.

1973

.

Behavior, mimetic songs and song dialects, and relationships of the parasitic indigobirds (Vidua) of Africa

.

Ornithological Monographs no. 11, American Ornithologists' Union, Washington, DC

.

Payne

,

R. B.

1977

.

The ecology of brood parasitism in birds

.

Annu. Rev. Ecol. Syst.

8

:

1

–

28

.

Payne

,

R. B.

1982

.

Species limits in the indigobirds (Ploceidae, Vidua) of West Africa: mouth mimicry, song mimicry, and description of new species

.

Misc. Pub. Mus. Zool. Univ. Mich.

no. 162

.

Payne

,

R. B.

1996

.

Field identification of the indigobirds

.

Bull. Afr. Bird Club

3

:

14

–

25

.

Payne

,

R. B.

1997a

. Avian brood parasitism. pp.

338

–

369

in

D. H.

Clayton

and

J.

Moore

, eds. Host‐parasite evolution: general principles and avian models.

Oxford Univ. Press

, Oxford, U.K..

Payne

,

R. B.

1997b

.

Field identification of the brood‐parasitic whydahs Vidua and cuckoo finch Anomalospiza

.

Bull. Afr. Bird Club

4

:

18

–

28

.

Payne

,

R. B.

1998a

.

Brood parasitism in birds: strangers in the nest

.

Bioscience

48

:

377

–

386

.

Payne

,

R. B.

1998b

.

A new species of firefinch Lagonosticta from northern Nigeria, and its association with the Jos Plateau indigobird Vidua maryae

.

Ibis

140

:

368

–

381

.

Payne

,

R. B.

2002

. Species accounts: family Viduidae and family Estrildidae (part). In

C. H.

Fry

and

S.

Kieth

, eds. The birds of Africa. Vol.

7

.

Academic Press

, New York. in press.

Payne

,

R. B.

, and

K.

Payne

.

1977

.

Social organization and mating success in local song populations of village indigobirds, Vidua chalybeata

.

Z. Tierpsychol.

45

:

113

–

173

.

Payne

,

R. B.

, and

L. L.

Payne

.

1994

.

Song mimicry and species status of the indigobirds Vidua: associations with quail‐finch Ortygospiza atricollis, goldbreast Amandava subflava and brown twinspot Clytospiza monteiri

.

Ibis

136

:

291

–

304

.

Payne

,

R. B.

, and

L. L.

Payne

.

1995

.

Song mimicry and association of brood‐parasitic indigobirds (Vidua) with Dybowski's twinspot (Euschistospiza dybowskii)

.

Auk

112

:

649

–

658

.

Payne

,

R. B.

, and

L. L.

Payne

.

2002

. Begging for parental care from another species: behavioural specialization and generalization in brood‐parasitic finches. pp.

429

–

450

in

J.

Wright

and

M. L.

Leonard

, eds. The evolution of begging: competition, cooperation and communication.

Kluwer

, Dordrecht, The Netherlands.

Payne

,

R. B.

,

L. L.

Payne

, and

M. E. D.

Nhlane

.

1992

.

Song mimicry and species status of the green widowfinch Vidua codringtoni

.

Ostrich

63

:

86

–

97

.

Payne

,

R. B.

,

L. L.

Payne

, and

J. L.

Woods

.

1998

.

Song learning in brood parasitic indigobirds Vidua chalybeata: song mimicry of the host species

.

Anim. Behav.

55

:

1537

–

1553

.

Payne

,

R. B.

,

L. L.

Payne

,

J. L.

Woods

, and

M. D.

Sorenson

.

2000

.

Imprinting, sex and the origin of parasite‐host species associations in female brood parasitic indigobirds Vidua chalybeata

.

Anim. Behav.

59

:

69

–

81

.

Payne

,

R. B.

,

J. L.

Woods

, and

L. L.

Payne

.

2001

.

Parental care in estrildid finches: experimental tests of a model of Vidua brood parasitism

.

Anim. Behav.

62

:

473

–

483

.

Payne

,

R. B.

,

K.

Hustler

,

R.

Stjernstedt

,

K. M.

Sefc

, and

M. D.

Sorenson

.

2002

.

Behavioural and genetic evidence of a recent population switch to a novel host species in brood parasitic indigobirds Vidua chalybeata

.

Ibis

144

: in press.

Posada

,

D.

, and

K. A.

Crandall

.

1998

.

MODELTEST: testing the model of DNA substitution

.

Bioinformatics

14

:

817

–

818

.

Restall

,

R.

1997

. Munias and mannikins.

Yale Univ. Press

, New Haven, CT.

Robert

,

M.

, and

G.

Sorci

.

1999

.

Rapid increase of host defence against brood parasites in a recently parasitized area: the case of village weavers in Hispaniola

.

Proc. R. Soc. Lond. B

266

:

941

–

946

.

Roberts

,

A.

1917

.

Parasitism amongst finches

.

Ann. Transvaal Mus.

5

:

259

–

262

.

Roberts

,

A.

1939

.

Notes on the eggs of parasitic birds in South Africa. Part 2

.

Ostrich

10

:

100

–

117

.

Rothstein

,

S. I.

1982

.

Mechanisms of avian egg recognition: Which egg parameters elicit response by rejecter species

Behav. Ecol. Sociobiol.

11

:

229

–

239

.

Rothstein

,

S. I.

1990

.

A model system for coevolution: avian brood parasitism

.

Annu. Rev. Ecol. Syst.

21

:

481

–

508

.

Rothstein

,

S. I.

2001

.

Relic behaviours, coevolution and the retention versus loss of host defences after episodes of avian brood parasitism

.

Anim. Behav.

61

:

95

–

107

.

Sharpe

,

R. B.

1888

. Catalogue of birds in the British Museum. Vol.

12

.

British Museum

, London.

Sharpe

,

R. B.

1890

. Catalogue of birds in the British Museum. Vol.

13

.

British Museum

, London.

Sheldon

,

F. H.

,

C. E.

Jones

, and

K. G.

McCracken

.

2000

.

Relative patterns and rates of evolution in heron nuclear and mitochondrial DNA

.

Mol. Biol. Evol.

17

:

437

–

450

.

Shelley

,

G. E.

1905

. The birds of Africa. Vol.

4

.

R. H. Porter

, London.

Sibley

,

C. G.

, and

J. E.

Ahlquist

.

1990

. Phylogeny and classification of birds, a study in molecular evolution.

Yale Univ. Press

, New Haven, CT.

Sibley

,

C. G.

, and

B. L.

Monroe

Jr.

1990

. Distribution and taxonomy of birds of the world.

Yale Univ. Press

, New Haven, CT.

Simmons

,

M. P.

, and

H.

Ochoterena

.

2000

.

Gaps as characters in sequence‐based phylogenetic analyses

.

Syst. Biol.

49

:

369

–

381

.

Soler

,

M.

,

J. J.

Soler

,

J. G.

Martinez

,

T.

Perez‐Contreras

, and

A. P.

Møller

.

1998

.

Micro‐evolutionary change and population dynamics of a brood parasite and its primary host: the intermittent arms race hypothesis

.

Oecologia

117

:

381

–

390

.

Sorenson

,

M. D.

1999

.

TreeRot

.

Ver. 2. Dept. of Biology, Boston Univ. Boston, MA

.

Sorenson

,

M. D.

, and

R. C.

Fleischer

.

1996

.

Multiple independent transpositions of mitochondrial DNA control region sequences to the nucleus

.

Proc. Nat. Acad. Sci.

93

:

15239

–

15243

.

Sorenson

,

M. D.

, and

T. W.

Quinn

.

1998

.

Numts: a challenge for avian systematics and population biology

.

Auk

115

:

214

–

221

.

Sorenson

,

M. D.

,

J. C.

Ast

,

D. E.

Dimcheff

,

T.

Yuri

, and

D. P.

Mindell

.

1999

.

Primers for a PCR‐based approach to mitochondrial genome sequencing in birds and other vertebrates

.

Mol. Phylogenet. Evol.

12

:

105

–

114

.

Steel

,

M. A.

,

A. C.

Cooper

, and

D.

Penny

.

1996

.

Confidence intervals for the divergence time of two clades

.

Syst. Biol.

45

:

127

–

134

.

Steyn

,

P.

1996

. Nesting birds, the breeding habits of southern African birds.

Fernwood Press

, Vlaeberg, South Africa.

Summers‐Smith

,

J. D.

1988

.

The sparrows, a study of the genus Passer

.

T. and A. D. Poyser, Calton, U.K

.

Swofford

,

D. L.

2000

. PAUP*: phylogenetic analysis using parsimony (*and other methods). Ver. 4.

Sinauer

, Sunderland, MA.

Swofford

,

D. L.

,

G. J.

Olsen

,

P. J.

Waddell

, and

D. M.

Hillis

.

1996

. Phylogenetic inference. pp.

407

–

514

in

D. M.

Hillis

,

C.

Moritz

, and

B. K.

Mable

, eds. Molecular systematics.

Sinauer

, Sunderland, MA.

Takasu

,

F.

1998

.

Why do all host species not show defense against avian brood parasitism: Evolutionary lag or equilibrium

Am. Nat.

151

:

193

–

205

.

Templeton

,

A. R.

1983

.

Phylogenetic inference from restriction endonuclease cleavage site maps with particular reference to the humans and apes

.

Evolution

37

:

221

–

244

.

Thompson

,

J. D.

,

D. G.

Higgins

, and

T. J.

Gibson

.

1994

.

Clustal W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, positions‐specific gap penalties and weight matrix choice

.

Nucleic Acids Res.

22

:

4673

–

4680

.

van Someren

,

V. G. L.

, and

G. R. C.

van Someren

.

1945

.

Evacuated weaver colonies and notes on the breeding ecology of Euodice cantans Gmelin and Amadina fasciata Gmelin

.

Ibis

87

:

33

–

44

.

van Tuinen

,

M.

,

C. G.

Sibley

, and

S. B.

Hedges

.

2000

.

The early history of modern birds inferred from DNA sequences of nuclear and mitochondrial ribosomal genes

.

Mol. Biol. Evol.

17

:

451

–

457

.

Vernon

,

C. J.

1964

.

The breeding of the cuckoo‐weaver (Anomalospiza imberbis (Cabanis)) in Southern Rhodesia

.

Ostrich

35

:

260

–

263

.

Vernon

,

C. J.

1987

.

Comments on some birds of Zimbabwe

.

Honeyguide

33

:

54

–

57

.

Wakeley

,

J.

1996

.

The excess of transitions among nucleotide substitutions: new methods of estimating transition bias underscore its significance

.

Trends Ecol. Evol.

11

:

158

–

163

.

Wernegreen

,

J. J.

, and

N. A.

Moran

.

1999

.

Evidence for genetic drift in endosymbionts (Buchnera): analyses of protein‐coding genes

.

Mol. Biol. Evol.

16

:

83

–

97

.

Wheeler

,

W. C.

1995

.

Sequence alignment, parameter sensitivity and the phylogenetic analysis of molecular data

.

Syst. Biol.

44

:

321

–

331

.

Wheeler

,

W. C.

1996

.

Optimization alignment: the end of multiple sequence alignment in phylogenetics

Cladistics

12

:

1

–

9

.

Wheeler

,

W. C.

1998

. Alignment characters, dynamic programming and heuristic solutions. pp.

243

–

251

in

R.

DeSalle

and

B.

Schierwater

, eds. Molecular approaches to ecology and evolution.

Birkháuser

, Berlin.

Williams

,

J. G.

, and

G. S.

Keith

.

1962

.

A contribution to our knowledge of the parasitic weaver, Anomalospiza imberbis

.

Bull. Brit. Ornithol. Club

82

:

141

–

142

.

Ziswiler

,

V.

1965

.

Zur Kenntnis des Samenöffnens und der Struktur des hörnernen Gaumens bei körnerfressenden Oscines

.

J. Ornithol.

106

:

1

–

48

.

Ziswiler

,

V.

1967

.

Vergleichend morphologische Untersuchungen am Verdauungstrakt körnerfressender Singvögel zur Abklärung ihrer systematischen Stellung

.

Zool. Jahrb. Syst.

94

:

427

–

520

.

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