Abstract
Female birds produce medullary bone to offset heightened calcium demand during egg shelling. The ephemeral nature of medullary bone hampers its observation in wild birds; thus, its phylogenetic distribution and ecophysiological correlates remain controversial. To address this, I candled a phylogenetically broad sample of bird skeletons spanning a large range of body sizes. I hypothesized that the presence of medullary bone would reduce or eliminate light penetrance through long bones. Candling suggested medullary bone was present in many adult female specimens, which was confirmed visually or with digital X-rays. This survey yielded the first unambiguous evidence of medullary bone in several taxa, including hummingbirds, tinamous, megapodes, and kiwi. The phylogenetic distribution of these new occurrences confirms that medullary bone is phylogenetically widespread among birds, including palaeognaths and passerines. The rapid assessment of complete skeletons supports a relationship between medullary bone and hematopoietic marrow, and establishes that the skeletal distribution of medullary bone varies taxonomically. Because the presence of medullary bone confirms sex, maturity, and reproductive status, its detection in specimens of known collection date can improve our understanding of life history and reproductive timing in wild birds, even decades later.
Zusammenfassung
Medullärer Knochen ist stammesgeschichtlich weit verbreitet, aber wo im Skelett er auftritt, variiert zwischen Taxa
Vogelweibchen produzieren medullären Knochen, um ihren erhöhten Kalziumbedarf während der Produktion von Eierschalen zu stillen. Die temporäre Natur dieses Knochengewebes erschwert seinen Nachweis in Wildvögeln, weshalb seine stammesgeschichtliche Verbreitung und ökophysiologischen Korrelate nach wie vor umstritten sind. Daher habe ich eine stammesgeschichtlich breite Stichprobe von Vogelskeletten unterschiedlicher Körpergröße durchleuchtet. Ich habe die Hypothese aufgestellt, dass das Vorhandensein medullären Knochens die Lichtdurchdringung von Röhrenknochen verringern oder ganz verhindern sollte. Das Durchleuchten deutete darauf hin, dass medullärer Knochen bei vielen Exemplaren adulter Weibchen vorhanden war, was dann visuell oder mit Hilfe digitaler Röntgenbilder bestätigt wurde. Diese Untersuchung lieferte den ersten eindeutigen Beleg für das Vorhandensein medullären Knochens in mehreren Taxa, einschließlich Kolibris, Steißhühnern, Großfußhühnern und Kiwis. Die stammesgeschichtliche Verbreitung dieser neuen Nachweise bestätigt, dass medullärer Knochen bei Vögeln stammesgeschichtlich weit verbreitet ist und bei Urkiefer- und Sperlingsvögeln vorkommt. Eine schnelle Prüfung vollständiger Skelette stützt einen Zusammenhang zwischen medullärem Knochen und blutbildendem Knochenmark und zeigt, dass die Verteilung medullären Knochens im Skelett zwischen Taxa variiert. Da das Vorkommen medullären Knochens Geschlecht, Geschlechtsreife und Fortpflanzungsstatus bestätigt, kann sein Nachweis in datierten Sammlungsexemplaren selbst mehrere Jahrzehnte später zu einem besseren Verständnis der Lebensgeschichte und des Fortpflanzungstimings von Wildvögeln führen.
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Acknowledgements
I thank Helen James and Christopher Milensky for access to USNM Bird Specimens, Sandra Raredon for access to and training on the USNM digital X-ray, and Matthew Brown and Chris Sagebiel for access to TMM specimens. Mary Schweitzer and Kevin Padian provided valuable discussion about the appearance and distribution of medullary bone. I thank Franz Bairlein, Daniel Field, and two anonymous reviews and for constructive comments that improved this manuscript. This work complies with the current laws of the country in which they were performed.
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Communicated by F. Bairlein.
Electronic supplementary material
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10336_2017_1514_MOESM1_ESM.pdf
An expanded discussion of methods and results, additional references for Fig. 3, and X-ray images for all 22 X-rayed specimens listed in Tables 1 and S1 (PDF 23971 kb)
10336_2017_1514_MOESM2_ESM.xlsx
Full dataset of specimens examined (includes additional notes, reproductive data, and natural history data) (XLSX 32 kb)
Video illustrating the candling method, using USNM 560556 (female Phalacrocorax auritus). Video is also available at https://youtu.be/XHkBk7JhKaA (MOV 251974 kb)
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Werning, S. Medullary bone is phylogenetically widespread and its skeletal distribution varies by taxon. J Ornithol 159, 527–543 (2018). https://doi.org/10.1007/s10336-017-1514-z
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DOI: https://doi.org/10.1007/s10336-017-1514-z