Frederick H. Sheldon



I am interested in the evolution and ecology of birds in Southeast Asia. Most of my recent work has been focused in two areas:

(1) Population genetics and phylogenetics of closely related species.

(2) The effect of forest disturbance and plantation development on native birds.  Personally, I work almost exclusively in Malaysian Borneo, but I have students and colleagues working on projects from China to Indonesia.  Our main goal is to understand forces that are related to the diversification of birds in the region.




Yale University:  Ph.D. Biology, 1986; M.S. Biology, 1981; B.S. Geology & Geophysics, 1974

Ph.D. Dissertation

"A Study of the Evolution and Phylogeny of Herons (Ardeidae) using DNA-DNA Hybridization."  Advisor:  C.G. Sibley


Selected Publications


Burner, R.C., A.R. Styring, and F.H. Sheldon. Faunal overlap produces low-to-mid elevation richness peak in birds on Bornean mountains. Journal of Tropical Ecology. In press.

2018 Campillo, L.C., C.H. Oliveros, F.H. Sheldon, and R.G. Moyle. Genomic data resolve gene tree discordance in spiderhunters (Nectariniidae, Arachnothera). Molecular Phylogenetics and Evolution 120:151-157. link
2018 Pegan, T.M., Gulson-Castillo, E.R., Biun, A., Byington, J.I., Moyle, R.G., Tuh, F.Y.Y., Wood, E.M., Winkler, D.W. and Sheldon, F.H. An assessment of avifauna in a recovering lowland forest at Kinabalu National Park, Malaysian Borneo. Raffles Bulletin of Zoology 66:110-131. link
2018 Shakya, S.B., T. Haryoko, R.C. Burner, D. Prawiradilaga, and F.H. Sheldon. Preliminary assessment of community composition and phylogeographic relationships of the birds of the Meratus Mountains, Southeastern Borneo, Indonesia. Bulletin of the British Ornithologists’ Club 138:45-66. link
2018 Styring, A.R., J. Unggang, A. Jukie, O. Tateh, N. Megam, and F.H. Sheldon. Bird community interactions between native forest fragments and Acacia mangium plantation in Borneo. Wilson Journal of Ornithology. In press.  
2017 Moyle, R.G., J.D. Manthey, P.A. Hosner, M.A. Rahman, M. Lakim, and F.H. Sheldon. A genome-wide assessment of stages of elevational parapatry in Bornean passerine birds reveals no introgression: implications for processes and patterns of speciation. PeerJ 5:e3335. link

Manthey, J.D., R.G. Moyle, D.F. Gawin, M.A. Rahman, M.F.S. Ramji, and F.H. Sheldon. Genomic phylogeography of the endemic Mountain Blackeye of Borneo (Chlorocharis emiliae): montane and lowland populations differ in patterns of Pleistocene diversification. Journal of Biogeography 44:2272–2283.


Lim, H.C., D.F. Gawin, S.B. Shakya, M.G. Harvey, M.A. Rahman, and F.H. Sheldon. Sundaland’s east-west rain forest population structure: variable manifestations in four polytypic bird species examined using RAD-Seq and plumage analyses. Journal of Biogeography 44: 2259–2271.

2017 Chua, V.L., B.T. Smith, M.A. Rahman, M. Lakim, D.M. Prawiradilaga, R.G. Moyle, and F.H. Sheldon. Evolutionary and ecological forces influencing population diversification in Bornean montane passerines. Molecular Phylogenetics and Evolution 113:139-149. link
2017 Reddy, S., R.T. Kimball, A. Pandey, P. Hosner, M.J. Braun, S.J. Hackett, K.L. Han, J. Harshman, C.J. Huddleston, S. Kingston, B.D. Marks, K.J. Miglia, W.A. Moore, F.H. Sheldon, T. Yuri, C.C. Witt,  E.L. Braun. Why do phylogenomic data sets yield conflicting trees? Data type influences the avian tree of life more than taxon sampling. Systematic Biology 66:857-879. link
2016 Burner, R.C., V.L. Chua, M.L. Brady, P. Van Els, P.O.M. Steinhoff, M.A. Rahman, and F.H. Sheldon. An ornithological survey of Gunung Mulu National Park, Sarawak, Malaysian Borneo. Wilson Journal of Ornithology 128:242-254. link

Cibois, A., and F.H. Sheldon. Comment on Jukema et al. “Geographic variation in morphometrics, molt, and migration suggests ongoing subspeciation in Pacific Golden-Plovers (Pluvialis fulva)”. The Auk: Advances in Ornithology 133:129-130.

2016 Styring, A.R., R. Ragai, and F.H. Sheldon. Foraging ecology and occurrence of seven sympatric babbler species (Aves: Timaliidae) in the lowland rainforest of Borneo and peninsular Malaysia. Current Zoology 62:345-355. link
2015 Sheldon, F.H. Gazetteer and site-based history of ornithology in Sabah, Malaysian Borneo. Occasional Papers of the Museum of Natural Science, Louisiana State University. link
2015 Chua, V. L., Q. Phillipps, H. C. Lim, S. S. Taylor, D. F. Gawin, M. A. Rahman, R. G. Moyle, and F. H. Sheldon. Phylogeography of three endemic birds of Maratua Island, a potential archive of Bornean biogeography. Raffles Bulletin of Zoology 63:259-269. link
2015 Lim, H.C., F. Zou, and F.H. Sheldon.  Genetic differentiation in two widespread, open-forest bird species of Southeast Asia (Copsychus saularis and Megalaima haemacephala): Insights from ecological niche modeling.  Current Zoology online. link
2015 Sheldon, F.H., H.C. Lim, and R.G. Moyle.  Return to the Malay Archipelago: the biogeography of Sundaic rainforest birds.  Journal of Ornithology online, DOI 10.1007/s10336-015-1188-3. link
2014 Gawin, D.F., M.A. Rahman, M.F.S. Ramji, B.T. Smith, H.C. Lim, R.G. Moyle, and F.H. Sheldon. Patterns of avian diversification in Borneo: the case of the endemic Mountain Black-eye (Chlorocharis emiliae). The Auk: Advances in Ornithology 131:86-99. link
2014 Jarvis, E.D., et al. Whole-genome analyses resolve early branches in the tree of life of modern birds. Science 346:1320-1331. link
2014 Lim, H.C., V.L. Chua, P.M. Benham, C.H. Oliveros, M.A. Rahman, R.G. Moyle, and F.H. Sheldon*. Divergence history of the Rufous-tailed Tailorbird (Orthotomus sericeus) of Sundaland: Implications for the biogeography of Palawan and the taxonomy of island species in general. Auk:Advances in Ornithology 131:629-642. link
2014 Sheldon, F.H., G. Davison, A. Wong, and R.G. Moyle.  Birds in peatswamp at Klias Forest Reserve and environs, Sabah, Malaysian Borneo.  Occasional Papers of the Museum of Natural Science, Louisiana State University 83:1-29 link
2014 Zhang, G., et al. Comparative genomics reveals insights into avian genome evolution and adaptation. Science 346:1311-1320. link
2013 Li, S., F. Zou, Q. Zhang, and F.H. Sheldon. Species richness and guild composition in rubber plantations compared to secondary forest on Hainan Island, China. Agroforestry Systems 87:1117-1128. link
2013 Mahood, S.P., A.J.I. John, J.C. Eames, C.H. Oliveros, R.G. Moyle, H. Chamnan, C.M. Poole, H. Nielsen, and F.H. Sheldon. 2013. A new species of lowland tailorbird (Passeriformes: Cisticolidae: Orthotomus) from the Mekong floodplain of Cambodia. Forktail 29:1-14. link
2013 Senior, M.J.M., K.C. Hamer, S.H. Bottrell, D.P. Edwards, T.M. Fayle, J.M. Lucey, P.J. Mayhew, R.J. Newton, K.S.H. Peh, F.H. Sheldon, C. Stewart, A.R. Styring, M.D. Thom, P. Woodcock, and J.K. Hill. Trait-dependent declines of species following conversion of rain forest to oil palm plantations. Biodiversity and Conservation 22:253-268. link
2013 Sheldon, F.H., C.E. Brown, M.A. Rahman, K.T. Guan, and R.G. Moyle. Ornithology of the Kelabit Highlands of Sarawak, Malaysia. Raffles Bulletin of Zoology. 61:1-27. link
2013 Yuri, T., R.T. Kimball, J. Harshman, R.C.K. Bowie, M.J. Braun, J. L. Chojnowski, S. Hackett, K.-L. Han, C.J. Huddleston, W.S. Moore, S. Reddy, F.H. Sheldon, D.W. Steadman, C.C. Witt, and E.L. Braun. Parsimony and model-based analyses of indels in avian nuclear genes reveal congruent and incongruent phylogenetic signals. Biology 2:419-444 link
2012 Dor, R., M.D. Carling, I.J. Lovette, F.H. Sheldon, and D.W. Winkler. Species trees for the tree swallows (Genus Tachycineta): an alternative phylogenetic hypothesis to the mitochondrial gene tree. Molecular Phylogenetics and Evolution 65:317-322. link
2012 Harris, J.B.C., D.L. Yong, F.H. Sheldon, A.J. Boyce, J.A. Eaton, H. Bernard, A. Biun, A. Langevin, T.E. Martin, and D. Wei. Using diverse data sources to detect elevational range changes of birds on Mount Kinabalu, Malaysian Borneo. Raffles Bulletin of Zoology Supplement 25:197-247. link
2012 Sheldon, F.H., C.H. Oliveros, S.S. Taylor, B. McKay, H.C. Lim, M.A. Rahman, H. Mays, and R.G. Moyle.  Molecular phylogeny and insular biogeography of the lowland tailorbirds of Southeast Asia (Cisticolidae: Orthotomus). Molecular Phylogenetics and Evolution 65:54-63. link
2012 Wu, Y., J.H. Huang, M. Zhang, S. Luo, Y. Zhang, F. Lei, F.H. Sheldon, and F. Zou. Genetic divergence and population demography of the Hainan endemic black-throated laughingthrush (Aves: Timaliidae, Garrulax chinensis monachus) and adjacent mainland subspecies. Molecular Phylogenetics and Evolution 65:482-489. link
2011 Braun, E.L., R.T. Kimball, K.-L. Han, N.R. Iuhasz, A.J. Bonilla, J.L. Chojnowski, J.V. Smith, R.C.K. Bowie, M J. Braun, S.J. Hackett, J. Harshman, C.J. Huddleston, B.D. Marks, K.J. Miglia, W.S. Moore, S. Reddy, F.H. Sheldon, C.C. Witt & T. Yuri. Homoplastic microinversions and the avian tree of life. BMC Evolutionary Biology 11:141. link
2011 Han, K.-L., E.L. Braun, R.T. Kimball, S. Reddy, R.C.K. Bowie, M.J. Braun, J.L. Chojnowski, S. Hackett, J. Harshman, C.J. Huddleston, B.D. Marks, K.J. Miglia, W.S. Moore, F.H. Sheldon, D.W. Steadman, C.C. Witt, and T. Yuri. Are transposable element insertions homoplasy free? An examination using the avian tree of life. Systematic Biology 60:375-386. link
2011 Lim, H.C., M.A. Rahman, S.L.H. Lim, R.G. Moyle, F.H. Sheldon. Revisiting Wallace’s haunt: coalescent simulations and comparative niche modeling reveal historical mechanisms that promoted avian population divergence in the Malay Archipelago. Evolution 65:321-334. link
2011 Lim, H.C., and F.H. Sheldon. Multilocus analysis of the evolutionary dynamics of rainforest bird populations in Southeast Asia. Molecular Ecology 20:3414-3438. link
2011 Moyle, R. G., S. S. Taylor, C. H. Oliveros, H. C. Lim, C. L. Haines, M. A. Rahman and F. H. Sheldon. Diversification of an endemic Southeast Asian genus: Phylogenetic relationships of the spiderhunters (Aves: Nectariniidae: Arachnothera). Auk 128:777-788. link
2011 Sheldon, F.H., and A.R. Styring. Bird diversity differs between industrial tree plantations on Borneo: implications for conservation planning. Raffles Bulletin of Zoology 59:269-283. link
2011 Styring, A.R., R. Ragai, J. Unggang, R. Stuebing, P.A. Hosner, and F.H. Sheldon. Bird community assembly in two Bornean tree plantations: effects of forest age and structure. Forest Ecology and Management 261:531-544. link

Amaral, F.S.R., F.H. Sheldon, A. Gamauf, E. Haring, M.J. Riesing, L.F. Silveira, and A. Wajntal.  Priority of Geranoaetus Kaup, 1844 over Tachytriorchis Kaup, 1844 (Aves: Accipitridae) based on the first reviser principle. Zootaxa 2534:67-68.

2010 Amaral, F.S.R., F.H. Sheldon, and A. Wajntal.  Towards an assessment of character interdependence in avian RNA phylogenetics: a general secondary structure model for the avian mitochondrial 16S rRNA.  Molecular Phylogenetics and Evolution 56:498-506. link

Dor, R., R.J. Safran, F.H. Sheldon, D.W. Winkler, and I.J. Lovette. Phylogeny of the genus Hirundo and the barn swallow subspecies complex. Molecular Phylogenetics and Evolution 56:409-418.

2010 Hosner, P.A., F.H. Sheldon, H.C. Lim, and R.G. Moyle. Phylogeny and biogeography of the Asian trogons (Aves: Trogoniformes) inferred from nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 57:1219-1215. link

Lim, H.C., F.H. Sheldon, and R.G. Moyle.  Extensive color polymorphism in the Southeast Asian oriental dwarf kingfisher Ceyx erithaca: a result of gene flow during population divergence?  Journal of Avian Biology 41:305-318.

2010 .  Lim, H.C., Zou, F., Taylor, S.S., Marks, B.D., Moyle, R.G., Voelker, G., and Sheldon, F.H. Phylogeny of magpie-robins and shamas (Aves: Turdidae: Copsychus and Trichixos): implications for island biogeography in Southeast Asia. Journal of Biogeography 37:1894-1906. link
2010 Lohman, D.J., K.K. Ingram, D.M. Prawiradilaga, K. Winker, F.H. Sheldon, R.G. Moyle, P.K.L. Ng, P.S. Ong, L.K. Wang, T.M. Braile, D. Astutic, and R. Meier. 2009. Cryptic genetic diversity in "widespread" Southeast Asian bird species suggests that Philippine avian endemism is gravely underestimated. Biological Conservation.  143: 1885-1890. link
2010 Sheldon, F.H., A.R. Styring, and P.A. Hosner. Bird species richness in a Bornean exotic tree plantation: a long-term perspective.  Biological Conservation 143:399-407. link
2009 Amaral, F.S.R., F.H. Sheldon, A. Gamauf, E. Haring, M.J. Riesing, L.F. Silveira, and A. Wajntal, A.  Patterns and processes of diversification in a widespread and ecologically diverse avian group, the buteonine hawks (Aves, Accipitridae). Molecular Phylogenetics and Evolution. 53:703-715. link
2009 Kimball, R.T., Braun, E.L., Barker, F.K., Bowie, R.C.K., Braun, M.J., Chojnowski, J.L., Hackett, S., Han, K.-L., Harshman, J., Heimer-Torres, V., Holznagel, W., Huddleston, C.J., Marks, B.D., Miglia, K.J., Moore, W.S., Reddy, S., Sheldon, F.H., Smith, J.V., Witt, C.C. & Yuri, T. A well-tested set of primers to amplify regions spread across the avian genome. Molecular Phylogenetics and Evolution 50:654-660. link
2009 Sheldon, F.H., H.C. Lim, J. Nais, M. Lakim, A. Tuuga, P. Malim, J. Majuakim, A. Lo, M. Schilthuizen, P.A. Hosner, R.G. Moyle.  Observations on the ecology, distribution, and biogeography of forest birds in Sabah, Malaysia.  Raffles Bulletin of Zoology 57:577-586. link
2009 Sheldon, F.H., D. J. Lohman, H.C. Lim, F. Zou, S.M. Goodman, D.M. Prawiradilaga, K. Winker, T.M. Braile, and R.G. Moyle. Phylogeography of the magpie-robin species complex (Aves: Turdidae: Copsychus) reveals a Philippine species, an interesting isolating barrier, and unusual dispersal patterns in the Indian Ocean and Southeast Asia. Journal of Biogeography 36:1070-1083. link
2008 Hackett, S., R.T. Kimball, S. Reddy, R.C.K. Bowie, E.L. Braun, M.J. Braun, J.L. Chojnowski, W.A. Cox, K.-L. Han, J. Harshman, C.J. Huddleston, B.D. Marks, K.J. Miglia, W.S. Moore, F.H. Sheldon, D.W. Steadman, C.C. Witt, and T. Yuri. A phylogenomic study of birds reveals their evolutionary history. Science 320:1763-1768. link
2008 Harshman, J., E.L. Braun, M.J. Braun, C.J. Huddleston, R.C.K. Bowie, J.L. Chojnowski, S. Hackett, K.-L. Han, R.T. Kimball, B.D. Marks, K.J. Miglia, W.S. Moore, S. Reddy, F.H. Sheldon, D.W. Steadman, S.J. Steppan, C.C. Witt, and T. Yuri. Phylogenomic evidence for multiple losses of flight in ratite birds. Proceedings of the National Academy of Sciences, USA 105:13462-13467. link
2008 Moyle, R.G., Hosner, P.A., Nais, J., Lakim, M., and Sheldon, F.H. Taxonomic status of the Kinabalu 'linchi' swiftlet. Bulletin of the British Ornithologists' Club 128:94-100. link
2008 Moyle, R.G., B. Slikas, L.A. Whittingham, D.W. Winkler, and F.H. Sheldon.  DNA sequence assessment of phylogenetic relationships among New World martins (Hirundinidae: Progne).  Wilson Journal of Ornithology 120:683-691. link
2007 Zou, F.S., H.C. Lim, B. Marks, R.G. Moyle, and F. H. Sheldon.  Molecular phylogenetic analysis of the Grey-cheeked Fulvetta (Alcippe morrisonia) of China and Indochina:  A case of remarkable genetic divergence in a “species”.  Molecular Phylogenetics and Evolution 44:165-174. link
2006 Moyle, R.G., J. Cracraft, M. Lakim, J. Nais, and F.H. Sheldon. Reconsideration of the phylogenetic relationships of the enigmatic Bornean Bristlehead (Pityriasis gymnocephala). Molecular Phylogenetics and Evolution 39:893-898. link
2005 Gill, F.B., B. Slikas, and F.H. Sheldon.  Phylogeny of titmice (Paridae):  II.  Species relationships based on sequences of the mitochondrial cytochrome-b gene.  Auk 122:121-143. link
2005 Moyle, R.G., M. Schilthuizen, M.A. Rahman, and F.H. Sheldon.  Molecular phylogenetic analysis of the White-crowned Forktail (Enicurus leschenaultia) in Borneo.  Journal of Avian Biology 36:96-101. link
2005 Sheldon, F.H., L.A. Whittingham, R.G. Moyle, B. Slikas, and D.W. Winkler. Phylogeny of swallows (Aves: Hirundinidae) estimated from nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 35:254-270. link