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NESTLING MOUTH Marklngs It '" "' of OLD WORLD FINCHES ESTLLU MIMICRY and COEVOLUTION of NESTING

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NESTLING MOUTH Marklngs It ' NESTLING MOUTH MARklNGS It '" "' OF OLD WORLD FINCHES ESTLLU MIMICRY AND COEVOLUTION OF NESTING r - .. ;.-; 5.i A&+.FINCHES .-. '4 AND THEIR VIDUA BROOD PARASITES - . , , . :.. - i ' -, ,' $*.$$>&.--: 7 -.: ',"L dt$=%>df;$..;,4;x.;b,?b;.:, ;.:. -, ! ,I Vt .., . k., . .,.-. , .is: 8, :. BY ERT B. PAYNE MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 194 Ann ntwi day, 2005 lSSN 0076-8405 PUBLICATIONS OF THE MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NO. 194 J. B. BLJR(.H,Editor JI.:NNIFERFBLMLEE, Assistcint Editor The publications of the Museum of Zoology, The University of Michigan, consist primarily of two series-the Mi.scel/aneous Pziblications and the Occa.siona1 Papers. Both series were founded by Dr. Bryant Walker, Mr. Bradshaw H. Swales, and Dr. W.W. Newcomb. Occasionally thc Museum publishes contributions outside of these series; beginning in 1990 thcsc arc titled Special Publications and arc numbered. All submitted manuscripts to any of the Museum's publications receive external review. The Occasional Papers, begun in 1913, serve as a medium for original studies based principally upon the collections in the Museum. They arc issued separately. When a sufficient number of pages has been printed to make a volume, a title page, table of contents, and an index are supplied to libraries and individuals on the mailing list for the series. The Miscellaneotls Pt~hlication.~,initiated in 1916, include monographic studies, papers on field and museum techniques, and other contributions not within the scope of the Occasional Papers, and are published separately. It is not intended that they be grouped into volurnes. Each number has a title page and, when necessary, a table of contents. A complete list of publications on Mammals, Birds, Reptiles and Amphibians, Fishes, Insects, Mollusks, and other topics is available. Address inquiries to Publications, Museum of Zoology, The University of Michigan, Ann Arbor, Michigan 48 109- 1079. RECENT MISCELLANEOUS PUBLICATIONS Lehtinen, R.M. (ed). 2004. Ecology and evolution of phytotelm-brccding anurans. Misc. PzrI11. MLIS.Zool., Univ. Michigan, No. 193, pp. i-iv, 1-73, 32 figs., 20 tables. Bailcy, R.M., W.C. Latta, & G.R. Smith. 2004. An atlas of Michigan fishes with keys and illustrations for their identification. Misc. Publ. Mzu. Zool., Univ. Michigun, No. 192, pp. i-iv, 1-2 15, 2 19 figs., 4 tablcs. Storer, R.W. 2002. The metazoan parasitc fauna of loons (Aves: Gaviiformcs), its relationship to the birds' evolutionary history and biology, and a comparison with the parasite fauna of grebes. Misc. Ptihl. MUS.Zoo/., Univ. Michigan, No. 191, pp. i-iv, 1 - 44, 3 figs., 7 tables. Albert, J.S. 2001. Species diversity and phylogenetic systematics of American knifefishes (Gymnotiformes, Teleostci). Misc. Publ. MLK.Zool., Uni~tMichigan, No. 190, pp. i-vi, 1-127, 50 figs. Nussbaum, R.A. & C.J. Raxworthy. 2000. Syste~naticrevision of the genus Pclroedtlm Giinther (Reptilia: Squamata: Gekkonidae), with the description of five new species. Misc. Ptibl. MLIS.Zool., Univ. Michigan, No. 189, pp. i-iv, 1-26, 12 figs., 7 tablcs. Storer, R.W. 2000. The metazoan parasite fauna of grcbes (Aves: Podicipediformes) and its relationship to the birds' biology. Misc. Pub/. Mza. Zool., Univ. Michigan, No. 188, pp. i-iv, 1-90, 9 figs., 7 tables. RECENT OCCASIONAL PAPERS Garrison, R.W., N. von Ellenrieder & M.F. O'Brien. 2003. An annotated list of the name-bearing types of species-group names in Odonata prcservcd in the University of Michigan Museum of Zoology. Occ. Pap. Mza. Zool., Univ. Michigan, No. 736, 73 pp. Ng, H.H. & J.S. Sparks. 2003. The ariid catfishes (Teleostei: Siluriformcs: Ariidae) of Madagascar, with the description of two new species. Occ. Pap. Mus. Zool., Univ. Michigan, No. 735, 21 pp., I l figs., 1 table. Ng, H.H. 2003. Aritls verruco.su.s, a new species of freshwater ariid catfish (Teleostei: Ariidac) from the Mekong River. Occ. Pap. M~rs.Zool., UIZ~IJ.Michigun, No. 734, 14 pp., 6 figs., 1 tablc. Norris, S.M. 2001. Osteology of thc southwestern dartcrs, Etheostoina (Oligocej~hal~~s)(Teleostei, Pcrcidae)-with comparison to other North American percid fishes. Occ. Pap. MLN.Zool., Univ. Michigun, No. 733,44 pp., 18 figs. Ng, H.M. & W.J. Rainboth. 200 1. A review of the sisorid catfish genus Oreoglanis (Silurifor~nes:Sisoridae) with descriptions of four new specics. Occ. Pup. MLIS.Zoo/., Univ. Michigan, No. 732, 34 pp., 13 figs., 3 tables. Collctte, B.B. 2001. 0psun~r.sdichrostomtrs, a new toadfish (Telcostei: Batrachoididae) from the westcrn Caribbean Sea and southern Gulf of Mexico. Occ. Pup. Mus. Zool., Univ. Michigun, No. 73 1, 16 pp., 5 figs. Fink, W.L. & A. Machado-Allison. 2001. Serra.salmtrs hastatus, a new species of piranha from Brazil, with comments on Ser- rnsalm~isalttivei and Serru.salmzis compresstis (Teleostci: Characiformes). Occ. Paj~.Mus. Zool., Univ. Michigan, No. 730, 18 pp., 16 figs. THE REGENTS OF THE UNIVERSITY David A. Brandon, Ann Arbor Rebecca McGowan, Ann Arbor S. Martin Taylor, Grosse Pointe Farms Laurence B. Dcitch, Blootnfield Hills Andrea Fischer Newman, Ann Arbor Katherine E. White, Ann Arbor Olivia P. Maynard, Goodrich Andrew C. Richner Mary Sue Coleman, ex oficio 0 Museum of Zoology, University of Michigan, 2005 Ann Arbor, Michigan 48109-1079, U.S.A. COVER ILLUSTRATION - Lagonosticta rara, Black-Bellied Firefinch MlSCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 194 Nestling Mouth Markings and Colors of Old World Finches Estrildidae: Mimicry and Coevolution of Nesting Finches and their Vidua Brood Parasites Robert B. Payne Museum of Zoology and Department of Ecology and Evolutionary Biology The University of Michigan Ann Arbor, Michigan 48109-1079, USA Ann Arbor, May, 2005 i NESTLING MOUTH MARKINGS AND COLORS OF OLD WORLD FINCHES ESTRILDIDAE: MIMICRY AND COEVOLUTION OF NESTING FINCHES AND THEIR VIDUA BROOD PARASITES CONTENTS PAGE ... Abstract .......................................................................................................................................................... nl Introduction ..................................................................................................................................................... 1 Methods .......................................................................................................................................................... 2 Systematics and sources of information .............................................................................................................. 2 Begging behaviors and parental care ................................................................................................................... 4 Morphology ..................................................................................................................................................... 4 Nestling development ....................................................................................................................................... 5 . Character scores and character ind~ces................................................................................................................ 6 Descriptions of estrildid finch species: nestling mouth markings and colors ............................................................ 13 Results ............................................................................................................................................................ 30 Phylogeny ........................................................................................................................................................ 30 Habitat ............................................................................................................................................................ 30 . Opportunistic nest parasltlsm by estr~ld~dfinches ................................................................................................. 3 1 Phylogeny, habitat, and character divergence ....................................................................................................... 3 1 Nestling mimicry by Vidua brood parasites ......................................................................................................... 32 Coevolution of nestling estrildids and their Vidua brood parasites .......................................................................... 34 Discussion ....................................................................................................................................................... 35 Behavioral constraints in parental care .............................................................................................................. 35 Mouth mimicry and coevolution in Vidua brood parasite and estrildid host ............................................................. 36 Coevolution between mimics and models in other animals ............................................................................... 38 Acknowledgments ............................................................................................................................................ 39 References ....................................................................................................................................................... 39 ABSTRACT Estrildid finches have remarkable colors and patterns in the mouths of their nestlings. Certain African estrildids rear young brood-parasitic finches, the indigobirds and whydahs Vidua, together
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