Cophylogenetic assessment of New World warblers (Parulidae) and their symbiotic feather mites (Proctophyllodidae)

Alix E. Matthews, Pavel B. Klimov, Heather C. Proctor, Ashley P.G. Dowling, Lizzie Diener, Stephen B. Hager, Jeffery L. Larkin, Douglas W. Raybuck, Cameron J. Fiss, Darin J. McNeil, Than J. Boves

Research output: Contribution to journalArticlepeer-review

20 Scopus citations


Host–symbiont relationships are ubiquitous in nature, yet evolutionary and ecological processes that shape these intricate associations are often poorly understood. All orders of birds engage in symbioses with feather mites, which are ectosymbiotic arthropods that spend their entire life on hosts. Due to their permanent obligatory association with hosts, limited dispersal and primarily vertical transmission, we hypothesized that the cospeciation between feather mites and hosts within one avian family (Parulidae) would be perfect (strict cospeciation). We assessed cophylogenetic patterns and tested for congruence between species in two confamiliar feather mite genera (Proctophyllodidae: Proctophyllodes, Amerodectes) found on 13 species of migratory warblers (and one other closely related migratory species) in the eastern United States. Based on COI sequence data, we found three Proctophyllodes lineages and six Amerodectes lineages. Distance- and event-based cophylogenetic analyses suggested different cophylogenetic trajectories of the two mite genera, and although some associations were significant, there was little overall evidence supporting strict cospeciation. Host switching is likely responsible for incongruent phylogenies. In one case, we documented prairie warblers Setophaga discolor harboring two mite species of the same genus. Most interestingly, we found strong evidence that host ecology may influence the likelihood of host switching occurring. For example, we documented relatively distantly related ground-nesting hosts (ovenbird Seiurus aurocapilla and Kentucky warbler Geothlypis formosa) sharing a single mite species, while other birds are shrub/canopy or cavity nesters. Overall, our results suggest that cospeciation is not the case for feather mites and parulid hosts at this fine phylogenetic scale, and raise the question if cospeciation applies for other symbiotic systems involving hosts that have complex life histories. We also provide preliminary evidence that incorporating host ecological traits into cophylogenetic analyses may be useful for understanding how symbiotic systems have evolved.

Original languageEnglish
Article numbere01580
JournalJournal of Avian Biology
Issue number3
StatePublished - Mar 2018

Bibliographical note

Funding Information:
Acknowledgements – We thank S. Mironov for helping to morphologically identify feather mites. We are extremely grateful for all who collected feathers or donated specimens for our collection. We also thank K. Richardson, A. Turnbull, S. Wilcox and C. Francois for their work as undergraduate technicians on this project. V. Ellis and E. Sari (Univ. Federal de Minas Gerais), A. Sweet (Univ. of Illinois at Urbana-Champaign) and J. Balbuena (Univ. de València) provided helpful advice on (co)phylogenetic analyses. J. and J. Gearhart graciously provided housing. The molecular work for this study was conducted in the Genomic Diversity Laboratory of the Dept of Ecology and Evolutionary Biology at the Univ. of Michigan. Funding – This work received support from the Arkansas State Univ. Dept of Biological Sciences, Arkansas Audubon Society Trust, the Univ. of Michigan Dept of Ecology and Evolutionary Biology, the Pennsylvania Game Commission, and the Natural Resources Conservation Service. PBK was supported by the Russian Science Foundation (project no. 16-14-10109) and the Russian Foundation for Basic Research (no. 15-04-0s5185-a). Author contributions – AEM, TJB, HCP and APGD conceived and designed the project. AEM, TJB, PBK, SBH and JLL received funding for the work. AEM, TJB, LD, SBH, JLL, DWR, CJF and DJM conducted field work. APGD contributed morphological and molecular analysis tools. HCP conducted laboratory work to morphologically identify mites. AEM and PBK performed molecular work and analyzed the data. AEM led the writing process, but all authors were involved in writing or constructively revising the manuscript. Conflicts of interest – The authors declare they have no conflict of interest. Permits – Capturing and handling of birds was conducted under United States Geological Survey federal banding permits no. 23877, no. 23277 and no. 23340, and was conducted in accordance with the guidelines of the Institutional Animal Care and Use Committee (IACUC) of Arkansas State Univ. (no. 63-8363) and Indiana Univ. of Pennsylvania (no. 14-1314). Carcasses collected were salvaged under state Scientific Permit no. NH11.0313 (Illinois Dept of Natural Resources) and federal Salvage Permit no. MB08907A-0 (US Fish and Wildlife Service).

Publisher Copyright:
© 2017 The Authors


  • Acari
  • birds
  • coevolution
  • cophylogenetic analysis
  • cospeciation
  • host–symbiont
  • symbiosis

ASJC Scopus subject areas

  • Ecology, Evolution, Behavior and Systematics
  • Animal Science and Zoology


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