1,2,3 = 2,3,4: a solution to the problem of the homology of the digits in the avian hand.

Persistent contradictions in well supported empirical findings usually point to important scientific problems and may even lead to exciting new insights. One of the most enduring problems in evolutionary biology is the apparent conflict between paleontological and embryological evidence regarding the homology of the digits in the avian hand (1, 2). We propose that this problem highlights an important feature of morphological change: namely, the possible dissociation between the developmental origin of a particular repeated element and its subsequent individualization into a fully functional character. We argue that, although comparative embryological evidence correctly identifies the homology of the primordial condensations in avians as CII, CIII, and CIV, subsequent anatomical differentiation reflects a frame shift in the developmental identities of the avian digit anlagen in later ontogeny such that CII becomes DI, CIII becomes DII, and CIV becomes DIII.

[1]  D. Duboule,et al.  Regulation of number and size of digits by posterior Hox genes: a dose-dependent mechanism with potential evolutionary implications. , 1997, Proceedings of the National Academy of Sciences of the United States of America.

[2]  P. Alberch,et al.  Size dependence during the development of the amphibian foot. Colchicine-induced digital loss and reduction. , 1983, Journal of embryology and experimental morphology.

[3]  Thomas R. Holtz,et al.  The phylogenetic position of the Tyrannosauridae: implications for theropod systematics , 1994, Journal of Paleontology.

[4]  J. Izpisúa-Belmonte,et al.  Molecular and cellular basis of pattern formation during vertebrate limb development. , 1999, Current topics in developmental biology.

[5]  P. Sereno,et al.  Dinosaurian precursors from the Middle Triassic of Argentina: Lagerpeton chanarensis , 1994 .

[6]  B. Hall,et al.  Divide, accumulate, differentiate: cell condensation in skeletal development revisited. , 2004, The International journal of developmental biology.

[7]  Arnold G. Kluge,et al.  AMNIOTE PHYLOGENY AND THE IMPORTANCE OF FOSSILS , 1988, Cladistics : the international journal of the Willi Hennig Society.

[8]  Günter P. Wagner,et al.  How can a character be developmentally constrained despite variation in developmental pathways? , 1993 .

[9]  P. Alberch,et al.  The development and homology of the chelonian carpus and tarsus , 1985, Journal of morphology.

[10]  John H. Ostrom,et al.  Archaeopteryx and the origin of birds , 1976 .

[11]  Pei-ji Chen,et al.  An exceptionally well-preserved theropod dinosaur from the Yixian Formation of China , 1998, Nature.

[12]  L. Jacobs,et al.  Aspects of vertebrate history: Essays in honor of Edwin Harris Colbert , 1980 .

[13]  Brian K. Hall,et al.  Homology: The hierarchical basis of comparative biology , 1994 .

[14]  C. Tabin,et al.  Molecular Models for Vertebrate Limb Development , 1997, Cell.

[15]  C. Nüsslein-Volhard,et al.  Mutations affecting segment number and polarity in Drosophila , 1980, Nature.

[16]  Craig Nelson,et al.  Hox genes and the evolution of vertebrate axial morphology. , 1995, Development.

[17]  P. Alberch,et al.  A morphogenetic approach to the origin and basic organization of the tetrapod limb , 1986 .

[18]  P. Sereno THE ORIGIN AND EVOLUTION OF DINOSAURS , 1997 .

[19]  K. Queiroz**,et al.  The Ontogenetic Method for Determining Character Polarity and its Relevance to Phylogenetic Systematics , 1985 .

[20]  P. Sereno The pectoral girdle and forelimb of the basal theropod Herrerasaurus ischigualastensis , 1994 .

[21]  A. Feduccia The origin and evolution of birds , 1996 .

[22]  P Chambon,et al.  Hoxa-13 and Hoxd-13 play a crucial role in the patterning of the limb autopod. , 1996, Development.

[23]  John H. Ostrom,et al.  The Origin of Birds , 1926 .

[24]  P. Alberch,et al.  Ontogeny of the limb skeleton in Alligator mississippiensis: Developmental invariance and change in the evolution of archosaur limbs , 1990, Journal of morphology.

[25]  D. Duboule,et al.  How to make a limb? , 1994, Science.

[26]  C. Tabin,et al.  Targeted misexpression of Hox-4.6 in the avian limb bud causes apparent homeotic transformations , 1992, Nature.

[27]  Ann C. Burke,et al.  Developmental Patterns and the Identification of Homologies in the Avian Hand , 1997 .

[28]  P. Chambon,et al.  Genetic interactions of Hox genes in limb development: learning from compound mutants. , 1997, Current opinion in genetics & development.

[29]  M. Norell,et al.  Two feathered dinosaurs from northeastern China , 1998, Nature.

[30]  C. Tabin,et al.  Why we have (only) five fingers per hand: hox genes and the evolution of paired limbs. , 1992, Development.

[31]  J. F. Meckel System der vergleichenden Anatomie , 1821 .

[32]  R. Narbaitz Vertebrate Limb and Somite Morphogenesis , 1979 .