title: Incorporating color into integrative taxonomy: analysis of the varied tit (Sittiparus varius) complex in East Asia.
abstract: Species designations are critically important scientific hypotheses that serve as the foundational units in a wide range of biological subdisciplines. A growing realization that some classes of data fail to delimit species under certain conditions has led to increasingly more integrative taxonomies, whereby species discovery and hypothesis testing are based on multiple kinds of data (e.g., morphological, molecular, behavioral, ecological, etc.). However, although most taxonomic descriptions have been based on morphology, some key morphological features, such as color, are rarely quantified and incorporated into integrative taxonomic studies. In this article, we applied a new method of ultraviolet digital photography to measure plumage variation in a color-variable avian species complex, the varied tit (Sittiparus varius). Plumage measurements corroborated species limits defined by morphometric, mitochondrial DNA, and nuclear DNA disjunctions and provided the only evidence for distinguishing two recently evolved species. Importantly, color quantification also provided a justification for lumping putative taxa with no evidence of evolutionary independence. Our revised taxonomy thus refines conservation units for listing and management and clarifies the primary units for evolutionary studies. Species tree analyses, which applied the newly delimited species as operational taxonomic units, revealed a robust phylogenetic hypothesis for the group that establishes a foundation for future biogeographic analyses. This study demonstrates how digital photography can be used to incorporate color character variation into integrative taxonomies, which should lead to more informed, more rigorous, and more accurate assessments of biodiversity. [Color, digital photography, integrative taxonomy, Sittiparus varius, species delimitation, varied tit.].
authors.affiliation: Department of Ornithology, American Museum of Natural History, 200 Central Park W, New York City, NY 10024, USA; Cincinnati Museum Center, Geier Collections and Research Building, 1301 Western Avenue, Cincinnati, OH 45203, USA; Medium Altitude Experimental Station, Endemic Species Research Institute, COA, ROC, #1 Min-Shen East Road, Chichi, Nantou County, 552, Taiwan; Department of Life Science, Liaoning University, No. 66 Chongshan Middle Road, Huanggu District, Shenyang 110036, China; Graduate School of Media and Governance, Keio University SFC, 5322 Endo, Fujisawa, Kanagawa 252-0882, Japan; and Department of Zoology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki 305-0005, Japan bmckay@amnh.org.
authors.forename: Bailey D
authors.initials: BD
authors.lastname: McKay
authors.affiliation: Department of Ornithology, American Museum of Natural History, 200 Central Park W, New York City, NY 10024, USA; Cincinnati Museum Center, Geier Collections and Research Building, 1301 Western Avenue, Cincinnati, OH 45203, USA; Medium Altitude Experimental Station, Endemic Species Research Institute, COA, ROC, #1 Min-Shen East Road, Chichi, Nantou County, 552, Taiwan; Department of Life Science, Liaoning University, No. 66 Chongshan Middle Road, Huanggu District, Shenyang 110036, China; Graduate School of Media and Governance, Keio University SFC, 5322 Endo, Fujisawa, Kanagawa 252-0882, Japan; and Department of Zoology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki 305-0005, Japan.
authors.forename: Herman L
authors.initials: HL
authors.lastname: Mays
authors.affiliation: Department of Ornithology, American Museum of Natural History, 200 Central Park W, New York City, NY 10024, USA; Cincinnati Museum Center, Geier Collections and Research Building, 1301 Western Avenue, Cincinnati, OH 45203, USA; Medium Altitude Experimental Station, Endemic Species Research Institute, COA, ROC, #1 Min-Shen East Road, Chichi, Nantou County, 552, Taiwan; Department of Life Science, Liaoning University, No. 66 Chongshan Middle Road, Huanggu District, Shenyang 110036, China; Graduate School of Media and Governance, Keio University SFC, 5322 Endo, Fujisawa, Kanagawa 252-0882, Japan; and Department of Zoology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki 305-0005, Japan.
authors.forename: Cheng-Te
authors.initials: CT
authors.lastname: Yao
authors.affiliation: Department of Ornithology, American Museum of Natural History, 200 Central Park W, New York City, NY 10024, USA; Cincinnati Museum Center, Geier Collections and Research Building, 1301 Western Avenue, Cincinnati, OH 45203, USA; Medium Altitude Experimental Station, Endemic Species Research Institute, COA, ROC, #1 Min-Shen East Road, Chichi, Nantou County, 552, Taiwan; Department of Life Science, Liaoning University, No. 66 Chongshan Middle Road, Huanggu District, Shenyang 110036, China; Graduate School of Media and Governance, Keio University SFC, 5322 Endo, Fujisawa, Kanagawa 252-0882, Japan; and Department of Zoology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki 305-0005, Japan.
authors.forename: Dongmei
authors.initials: D
authors.lastname: Wan
authors.affiliation: Department of Ornithology, American Museum of Natural History, 200 Central Park W, New York City, NY 10024, USA; Cincinnati Museum Center, Geier Collections and Research Building, 1301 Western Avenue, Cincinnati, OH 45203, USA; Medium Altitude Experimental Station, Endemic Species Research Institute, COA, ROC, #1 Min-Shen East Road, Chichi, Nantou County, 552, Taiwan; Department of Life Science, Liaoning University, No. 66 Chongshan Middle Road, Huanggu District, Shenyang 110036, China; Graduate School of Media and Governance, Keio University SFC, 5322 Endo, Fujisawa, Kanagawa 252-0882, Japan; and Department of Zoology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki 305-0005, Japan.
authors.forename: Hiroyoshi
authors.initials: H
authors.lastname: Higuchi
authors.affiliation: Department of Ornithology, American Museum of Natural History, 200 Central Park W, New York City, NY 10024, USA; Cincinnati Museum Center, Geier Collections and Research Building, 1301 Western Avenue, Cincinnati, OH 45203, USA; Medium Altitude Experimental Station, Endemic Species Research Institute, COA, ROC, #1 Min-Shen East Road, Chichi, Nantou County, 552, Taiwan; Department of Life Science, Liaoning University, No. 66 Chongshan Middle Road, Huanggu District, Shenyang 110036, China; Graduate School of Media and Governance, Keio University SFC, 5322 Endo, Fujisawa, Kanagawa 252-0882, Japan; and Department of Zoology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki 305-0005, Japan.
authors.forename: Isao
authors.initials: I
authors.lastname: Nishiumi
mesh_terms: D000818:Animals; D049628:Body Size; D002965:Classification; D004272:DNA, Mitochondrial / Q000235:genetics; D005202:Asia, Eastern; D005241:Feathers / Q000737:chemistry*; D008969:Molecular Sequence Data; D046109:Passeriformes / Q000033:anatomy & histology / Q000145:classification* / Q000235:genetics / Q000502:physiology*; D010802:Phylogeny; D010858:Pigmentation / Q000502:physiology*; D013045:Species Specificity
chemical_list: D004272:DNA, Mitochondrial