title: Evolution of body morphology and beak shape revealed by a morphometric analysis of 14 Paridae species.
abstract: BACKGROUND
  Morphological characters of birds reflect their adaptive evolution and ecological requirements and are also relevant to phylogenetic relationships within a group of related species. The tits (Paridae) are known to be outwardly homogeneous in shape, with one aberrant member, the Ground Tit (Pseudopodoces humilis), which is quite different from its relatives in both body morphology and beak shape. We combined traditional measurements and geometric morphometrics to quantify the variation in body morphology and beak shape of 14 Paridae species distributed in China. Based on these results, we sought to assess the contribution of phylogeny, altitude and species interactions to the evolution of morphological traits.
  
  
  RESULTS
  The basic features for discriminating among the 14 species studied here were overall body size, the ratio of body and tail length to culmen and tarsus length, and beak shape (long/slender/pointy vs. short/robust/blunt). These dimensions clearly separate Ps. humilis and Melanochlora sultanea from the other species in shape space. Body length and PC3 of beak shape (round outline vs. straight outline) show significant phylogenetic signals. Across 14 species, altitude is related to tarsus, culmen length and PC1 of beak shape. Within Parus major, altitude is related to body weight, body length, culmen length and PC1 of body morphology. Morphological distances and geographic distances among species are positively correlated.
  
  
  CONCLUSIONS
  The body morphology of Paridae species shows extensive evolutionary changes, while their beak has mainly evolved along the long/slender/pointy vs. short/robust/blunt dimension. Only body length and beak curvature show a phylogenetic signal. Altitude correlates with multiple traits both across and within species, suggesting that altitude is an important factor in promoting morphological divergence. The deviant appearance of Ps. humilis corresponds to its foraging and feeding adaptations to high-altitude steppe habitats. Our results also show a higher level of morphological divergence with greater difference in distribution ranges among the Paridae species involved in this study.
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, 100101 China ; University of Chinese Academy of Sciences, Beijing, 100049 China.
authors.forename: Shimiao
authors.initials: S
authors.lastname: Shao
authors.affiliation: Guangdong Entomological Institute, Chinese Academy of Sciences, Guangdong, 510260 China.
authors.forename: Qing
authors.initials: Q
authors.lastname: Quan
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, 100101 China ; University of Chinese Academy of Sciences, Beijing, 100049 China.
authors.forename: Tianlong
authors.initials: T
authors.lastname: Cai
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, 100101 China.
authors.forename: Gang
authors.initials: G
authors.lastname: Song
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, 100101 China.
authors.forename: Yanhua
authors.initials: Y
authors.lastname: Qu
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, 100101 China.
authors.forename: Fumin
authors.initials: F
authors.lastname: Lei
keywords: Altitude; Distribution overlap; Geometric morphometrics; Morphology; Paridae; Phylogenetic relationship