{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T21:54:31Z","timestamp":1791237271371,"version":"4.1.0"},"reference-count":62,"publisher":"Wildlife Disease Association","issue":"2","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Journal of Wildlife Diseases"],"published-print":{"date-parts":[[2026,4,20]]},"abstract":"<jats:p>Birds are host to four orders of parasitic insects: flies (Diptera), lice (Psocodea: Phthiraptera), true bugs (Hemiptera), and fleas (Siphonaptera). These parasites can lower host fitness and thus require birds to invest in defenses to minimize fitness losses. In Puerto Rico, virulent parasites, such as Philornis spp. flies, threaten endangered species and subspecies of birds. Philornis larvae are hematophagous and burrow beneath the skin of both nestling and adult birds. Unfortunately, little is known regarding the prevalence and intensity of subcutaneous Philornis flies of adult birds in Puerto Rico. Here, we inspected 430 birds, representing 38 species, for Philornis infestation. We found Philornis larvae on only four birds, despite previous studies reporting high Philornis prevalence on adults of a single Puerto Rican species, Pearly-eyed Thrashers (Margarops fuscatus). Our results are consistent with other studies suggesting that the prevalence and intensity of Philornis is low in most adult and fledgling birds, compared to nestlings. Our data suggest that Philornis infestation is not a major threat to adult birds in Puerto Rico. In contrast, we show that parasitic lice are relatively common on Puerto Rican birds. Of 309 birds dusted for ectoparasites, 163 (53%) were infested with lice. None of the 309 birds were infested with parasitic fleas or bugs. Our results confirm other recent work showing relatively high prevalence and intensity of lice on birds in humid regions of the world.<\/jats:p>","DOI":"10.7589\/jwd-d-25-00151","type":"journal-article","created":{"date-parts":[[2026,4,20]],"date-time":"2026-04-20T05:58:07Z","timestamp":1776664687000},"page":"407-420","source":"Crossref","is-referenced-by-count":0,"title":["PREVALENCE AND INTENSITY OF PARASITIC INSECTS ON PUERTO RICAN BIRDS"],"prefix":"10.7589","volume":"62","author":[{"given":"Matthew M.","family":"Waller","sequence":"first","affiliation":[{"name":"School of Biological Sciences, University of Utah, 257 South 1400 East, Salt Lake City, Utah 84102, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicole A.","family":"Amedee","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Utah, 257 South 1400 East, Salt Lake City, Utah 84102, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hannah M.","family":"Warr","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Utah, 257 South 1400 East, Salt Lake City, Utah 84102, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sarah E.","family":"Bush","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Utah, 257 South 1400 East, Salt Lake City, Utah 84102, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dale H.","family":"Clayton","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Utah, 257 South 1400 East, Salt Lake City, Utah 84102, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4415","reference":[{"key":"30","doi-asserted-by":"crossref","unstructured":"MJ. Lehane,  \"The biology of blood-sucking in insects.\"  321 (2005)","DOI":"10.1017\/CBO9780511610493"},{"key":"5","doi-asserted-by":"publisher","unstructured":"\"AviList: the global avian checklist, v2025.\"  (2025)","DOI":"10.2173\/avilist.v2025"},{"key":"1","doi-asserted-by":"crossref","unstructured":"V Amrhein, B Scaar, M Baumann, et al.  \"Methods Ecol Evol.\"  3, \"Estimating adult sex ratios from bird mist netting data.\" 713 (2012)","DOI":"10.1111\/j.2041-210X.2012.00207.x"},{"key":"2","doi-asserted-by":"crossref","unstructured":"WJ. Arendt,  \"Auk.\"  102, \"Philornis ectoparasitism of pearly-eyed thrashers. II. Effects on adults and reproduction.\" 281 (1985)","DOI":"10.2307\/4086770"},{"key":"3","unstructured":"WJ. Arendt,  \"Ornitol Neotrop.\"  11, \"Impact of nest predators, competitors, and ectoparasites on pearly-eyed thrashers, with comments on the potential implications for Puerto Rican parrot recovery.\" 13 (2000)"},{"key":"4","doi-asserted-by":"crossref","unstructured":"W. Arendt,  \"Adaptions of an avian supertramp: Distribution, ecology, and life history of the pearly-eyed thrasher (Margarops fuscatus). General Technical Report 27.\"  404 (2006)","DOI":"10.2737\/IITF-GTR-27"},{"key":"6","doi-asserted-by":"crossref","unstructured":"CM. Bartlett,  \"J Parasitol.\"  79, \"Lice (Amblycera and Ischnocera) as vectors of Eulimdana spp. (Nematoda: Filarioidea) in charadriiform birds and the necessity of short reproductive periods in adult worms.\" 85 (1993)","DOI":"10.2307\/3283282"},{"key":"7","doi-asserted-by":"crossref","unstructured":"DT Booth, DH Clayton, BA Block,  \"Proc R Soc Lond B.\"  253, \"Experimental demonstration of the energetic cost of parasitism, in free-ranging hosts.\" 125 (1993)","DOI":"10.1098\/rspb.1993.0091"},{"key":"9","doi-asserted-by":"crossref","unstructured":"SE Bush, MM Waller, KM Davis, et al.  \"Ecol Evol.\"  14, \"Birds in arid regions have depauperate louse communities: Climate change implications?\" e70280 (2024)","DOI":"10.1002\/ece3.70280"},{"key":"10","doi-asserted-by":"crossref","unstructured":"SE Bush, MM Waller, JM Herman, et al.  \"Anim Behav.\"  201, \"Birds groom more in regions with higher parasite pressure: A comparison of temperate and subtropical American kestrels.\" 125 (2023)","DOI":"10.1016\/j.anbehav.2023.04.015"},{"key":"11","doi-asserted-by":"crossref","unstructured":"H Cadena-Ortiz, M Quiroga, E Bonaccorso,  \"Rev Mex Biodivers.\"  93, \"First report of the Philornis torquans complex in adult birds in continental Ecuador: Is parasitism by Philornis being underestimated?\" e934080 (2022)","DOI":"10.22201\/ib.20078706e.2022.93.4080"},{"key":"12","doi-asserted-by":"crossref","unstructured":"J Castro-Prieto, JM Wunderle, J Salguero-Far\u00eda, et al.  \"The Puerto Rico breeding bird atlas. General Technical report IITF-53.\"  311 (2021)","DOI":"10.2737\/IITF-GTR-53"},{"key":"16","doi-asserted-by":"crossref","unstructured":"DH Clayton  and DM Drown  \"J Parasitol.\"  87, \"Critical evaluation of five methods for quantifying chewing lice (Insecta: Phthiraptera)\" 1291 (2001)","DOI":"10.2307\/3285290"},{"key":"17","doi-asserted-by":"crossref","unstructured":"DH Clayton  and RD Price  \"Ann Entomol Soc Am.\"  92, \"Taxonomy of New World Columbicola (Phthiraptera: Philopteridae) from the Columbiformes (Aves), with descriptions of five new species.\" 675 (1999)","DOI":"10.1093\/aesa\/92.5.675"},{"key":"18","unstructured":"MS. Couri,  \"Myiasis in man and animals in the neotropical region: Bibliographic database.\"  \"Myiasis caused by obligatory parasites. Ia. Philornis Meinert (Muscidae).\" 51 (1999)"},{"key":"19","doi-asserted-by":"crossref","unstructured":"PF Cuervo, A Percara, L Monje, et al.  \"Med Vet Entomol.\"  35, \"Environmental variables determining the distribution of an avian parasite: The case of the Philornis torquans complex in South America.\" 284 (2021)","DOI":"10.1111\/mve.12488"},{"key":"20","doi-asserted-by":"crossref","unstructured":"CA Delannoy  and A Cruz  \"Bird\u2013parasite interactions: Ecology, evolution and behaviour.\"  \"Philornis parasitism and nestling survival of the Puerto Rican sharp-shinned hawk.\" 93 (1991)","DOI":"10.1093\/oso\/9780198577386.003.0005"},{"key":"21","doi-asserted-by":"crossref","unstructured":"MG Frixione, SE Bush, DH Clayton,  \"J Wildl Dis.\"  6, \"High ectoparasite loads of tropical birds: Chewing lice on Puerto Rican American kestrels (Falco sparverius).\" 241 (2025)","DOI":"10.7589\/JWD-D-24-00100"},{"key":"23","doi-asserted-by":"crossref","unstructured":"DR Gustafsson  and SE Bush  \"Zootaxa.\"  4313, \"Morphological revision of the hyperdiverse Brueelia-complex (Insecta: Phthiraptera: Ischnocera: Philopteridae) with new taxa, checklists and generic key.\" 1 (2017)","DOI":"10.11646\/zootaxa.4313.1.1"},{"key":"24","doi-asserted-by":"crossref","unstructured":"S Kolencik, O Sychra, KP Johnson, et al.  \"Insect Syst Divers.\"  8, \"The parasitic louse genus Myrsidea (Amblycera: Menoponidae): A comprehensive review and world checklist.\" 1 (2024)","DOI":"10.1093\/isd\/ixae007"},{"key":"25","doi-asserted-by":"crossref","unstructured":"JAH Koop  and DH Clayton  \"J Field Ornithol.\"  84, \"Evaluation of two methods for quantifying passeriform lice.\" 210 (2013)","DOI":"10.1111\/jofo.12020"},{"key":"26","doi-asserted-by":"crossref","unstructured":"JAH Koop, SK Huber, SM Laverty, et al.  \"PLoS One.\"  6, \"Experimental demonstration of the fitness consequences of an introduced parasite of Darwin's finches.\" e19706 (2011)","DOI":"10.1371\/journal.pone.0019706"},{"key":"28","doi-asserted-by":"crossref","unstructured":"JB. LaPergola,  \"Ibis.\"  165, \"Life-stage and sex influence Philornis ectoparasitism in a Neotropical woodpecker Melanerpes striatus with essential male parental care.\" 1235 (2023)","DOI":"10.1111\/ibi.13221"},{"key":"29","doi-asserted-by":"crossref","unstructured":"JB LaPergola, CJ Proctor, H Almonte, et al.  \"J Caribb Ornithol.\"  36, \"Mortality of an adult Antillean palm swift (Tachornis phoenicobia) associated with Philornis sp. parasitism in the Sierra De Bahoruco, Dominican Republic.\" 36 (2023)","DOI":"10.55431\/jco.2023.36.36-44"},{"key":"32","doi-asserted-by":"crossref","unstructured":"DE Manzoli, MJ Saravia-Pietropaolo, LR Antoniazzi, et al.  \"Int J Parasitol.\"  48, \"Contrasting consequences of different defence strategies in a natural multihost\u2013parasite system.\" 445 (2018)","DOI":"10.1016\/j.ijpara.2017.11.001"},{"key":"34","unstructured":"AG. Marshall,  \"The ecology of ectoparasitic insects.\"  459 (1981)"},{"key":"35","doi-asserted-by":"crossref","unstructured":"CJW McClure, HC Weaver, M Murillo, et al.  \"J Raptor Res.\"  57, \"Break even points in nest management of an endangered island endemic raptor.\" 44 (2023)","DOI":"10.3356\/JRR-22-39"},{"key":"37","doi-asserted-by":"crossref","unstructured":"SM McNew, GB Goodman, JR Y\u00e9pez, et al.  \"Oecologia.\"  192, \"Parasitism by an invasive nest fly reduces future reproduction in Gal\u00e1pagos mockingbirds.\" 363 (2020)","DOI":"10.1007\/s00442-019-04582-y"},{"key":"38","doi-asserted-by":"crossref","unstructured":"AP M\u00f8ller, E Arriero, E Lobato, et al.  \"Biol Rev.\"  84, \"A meta-analysis of parasite virulence in nestling birds.\" 567 (2009)","DOI":"10.1111\/j.1469-185X.2009.00087.x"},{"key":"39","doi-asserted-by":"crossref","unstructured":"JA O'Connor, J Robertson, S Kleindorfer,  \"Ethology.\"  120, \"Darwin's finch begging intensity does not honestly signal need in parasitised nests.\" 228 (2014)","DOI":"10.1111\/eth.12196"},{"key":"40","doi-asserted-by":"crossref","unstructured":"JP Owen, AC Nelson, DH Clayton,  \"Trends Parasitol.\"  26, \"Ecological immunology of bird\u2013ectoparasite systems.\" 530 (2010)","DOI":"10.1016\/j.pt.2010.06.005"},{"key":"41","unstructured":"RA P\u00e9rez-Rivera  and J Collazo Algar\u00edn  \"Science-Ciencia.\"  4, \"Aves en Puerto Rico parasitadas por la larva de la mosca del g\u00e9nero Neomusca.\" 50 (1976)"},{"key":"43","unstructured":"RD Price, DH Clayton, RA Hellenthal,  \"J Med Entomol.\"  36, \"Taxonomic review of Physconelloides (Phthiraptera: Philopteridae) from the Columbiformes (Aves), including descriptions of three new species.\" 195 (1999)"},{"key":"45","unstructured":"P. Pyle,  \"Identification guide to North American birds, part I.\"  689 (2022)"},{"key":"46","doi-asserted-by":"crossref","unstructured":"MA Quiroga, LD Monje, JP Arrabal, et al.  \"Rev Mex Biodivers.\"  87, \"New molecular data on subcutaneous Philornis (Diptera: Muscidae) from southern South America suggests the existence of a species complex.\" 1383 (2016)","DOI":"10.1016\/j.rmb.2016.10.018"},{"key":"47","doi-asserted-by":"crossref","unstructured":"MA Quiroga, TI Hayes, CD Hayes, et al.  \"Parasitol Res.\"  199, \"More than just nestlings: Incidence of subcutaneous Philornis (Diptera: Muscidae) nest flies in adult birds.\" 2337 (2020)","DOI":"10.1007\/s00436-020-06696-2"},{"key":"48","unstructured":"MT. Rivera Irizarry,  \"Impacto del parasitismo por Philornis (Neomusca) angustifrons (Loew) (Diptera: Muscidae) en el crecimiento, desarrollo y supervivencia de los pichones del Zorzal Pardo, Margarops fuscatus (Vieillot) (Passeriformes: Mimidae) en Mayag\u00fcez, Puerto Rico.\"  (1991)"},{"key":"49","doi-asserted-by":"crossref","unstructured":"FE Rheindt, PF Donald, DB Donsker, et al.  \"Biodivers Conserv.\"  34, \"AviList: A unified global bird checklist.\" 3359 (2025)","DOI":"10.1007\/s10531-025-03120-y"},{"key":"50","unstructured":"M Rothschild  and T Clay  \"Fleas, flukes and cuckoos. A study of bird parasites.\"  370 (1952)"},{"key":"51","doi-asserted-by":"crossref","unstructured":"SC. Schubert,  \"J Caribb Ornithol.\"  31, \"Nesting of the Puerto Rican emerald (Chlorostilbon maugaeus) and a record of nestling parasitism by botflies (Philornis sp.).\" 77 (2018)","DOI":"10.55431\/jco.2018.31.77-82"},{"key":"52","doi-asserted-by":"crossref","unstructured":"WS Seegar, EL Schiller, WJL Sladen, et al.  \"Science.\"  194, \"A mallophaga, Trinton anserium, as a cyclodevelopmental vector for a heartworm parasite of waterfowl.\" 739 (1976)","DOI":"10.1126\/science.982042"},{"key":"53","unstructured":"NFR Snyder, JW Wiley, CB Kepler,  \"The parrots of Luquillo: Natural history and conservation of the Puerto Rican parrot.\"  384 (1987)"},{"key":"54","doi-asserted-by":"crossref","unstructured":"O Sychra, S Kolencik, I Papousek, et al.  \"Arthropod Syst Phylogeny.\"  79, \"Myrsidea quadrifasciata (Phthiraptera: Amblycera) \u2013 a unique host generalist among highly host-specific chewing lice.\" 379 (2021)","DOI":"10.3897\/asp.79.e63975"},{"key":"55","unstructured":"D. Texeira,  \"Myiasis in man and animals in the neotropical region: Bibliographic database.\"  \"Myasis caused by obligatory parasites, Ib. General observations on the biology of species of genus Philornis meinert.\" 71 (1999)"},{"key":"56","doi-asserted-by":"crossref","unstructured":"LS Uhazy  and WJ Arendt  \"J Wildl Dis.\"  22, \"Pathogenesis associated with philornid myiasis (Diptera: Muscidae) on nestling pearly-eyed thrashers (Aves: Mimidae) in the Luquillo Rain Forest, Puerto Rico.\" 224 (1986)","DOI":"10.7589\/0090-3558-22.2.224"},{"key":"57","unstructured":"\"Recovery plan for the Puerto Rican parrot (Amazona vittata).\"  75pp (2009)"},{"key":"58","unstructured":"BA Walther  and DH Clayton  \"J Field Ornithol.\"  68, \"Dust-ruffling: A simple method for quantifying ectoparasite loads of live birds.\" 509 (1997)"},{"key":"60","unstructured":"NS. Williams,  \"J Kans Entomol Soc.\"  52, \"The Picicola (Mallophaga: Philopteridae) of the Passeriformes (Aves).\" 633 (1979)"},{"key":"61","unstructured":"JS. Wiseman,  \"J Kans Entomol Soc.\"  36, \"A new species of Osborniella (Mallophaga) from the groove-billed ani.\" 101 (1963)"},{"key":"62","doi-asserted-by":"crossref","unstructured":"GN. Wolcott,  \"J Agric Univ Puerto Rico.\"  32, \"The insects of Puerto Rico.\" 417 (1948)","DOI":"10.46429\/jaupr.v32i3.13612"},{"key":"33","doi-asserted-by":"crossref","unstructured":"DE Manzoli, MJ Saravia-Pietropaolo, SI Arce, et al.  \"Int J Parasitol.\"  51, \"Specialist by preference, generalist by need: Availability of quality hosts drives parasite choice in a natural multihost\u2013parasite system.\" 527 (2021)","DOI":"10.1016\/j.ijpara.2020.12.003"},{"key":"14","doi-asserted-by":"crossref","unstructured":"DH Clayton, RJ Adams, SE Bush,  \"Parasitic diseases of wild birds.\"  \"Phthiraptera, the chewing lice.\" 515 (2008)","DOI":"10.1002\/9780813804620.ch29"},{"key":"15","doi-asserted-by":"crossref","unstructured":"DH Clayton, SE Bush, KP Johnson,  \"Coevolution of life on hosts: Integrating ecology and history.\"  294 (2015)","DOI":"10.7208\/chicago\/9780226302300.001.0001"},{"key":"27","doi-asserted-by":"crossref","unstructured":"JAH Koop, JP Owen, SA Knutie, et al.  \"Ecol Evol.\"  3, \"Experimental demonstration of a parasite-induced immune response in wild birds: Darwin's finches and introduced nest flies.\" 2514 (2013)","DOI":"10.1002\/ece3.651"},{"key":"31","doi-asserted-by":"crossref","unstructured":"DE Manzoli, LR Antoniazzi, MJ Saravia, et al.  \"PLoS One.\"  8, \"Multi-level determinants of parasitic fly infection in forest passerines.\" e67104 (2013)","DOI":"10.1371\/journal.pone.0067104"},{"key":"36","doi-asserted-by":"crossref","unstructured":"SM McNew  and DH Clayton  \"Annu Rev Entomol.\"  63, \"Alien invasion: Biology of Philornis flies highlighting Philornis downsi, an introduced parasite of Gal\u00e1pagos Birds.\" 369 (2018)","DOI":"10.1146\/annurev-ento-020117-043103"},{"key":"13","doi-asserted-by":"crossref","unstructured":"DH. Clayton,  \"Am Zool.\"  30, \"Mate choice in experimentally parasitized rock doves: lousy males lose.\" 251 (1990)","DOI":"10.1093\/icb\/30.2.251"},{"key":"59","doi-asserted-by":"crossref","unstructured":"M Wikelski, J Foufopoulos, H Vargas, et al.  \"Ecol Soc.\"  9, \"Gal\u00e1pagos birds and diseases: invasive pathogens as threats for island species.\" 5 (2004)","DOI":"10.5751\/ES-00605-090105"},{"key":"8","doi-asserted-by":"crossref","unstructured":"AO Bush, KD Lafferty, JM Lotz, et al.  \"J Parasitol.\"  83, \"Parasitology meets ecology on its own terms: Margolis et al. revisited.\" 575 (1997)","DOI":"10.2307\/3284227"},{"key":"44","unstructured":"RD Price, RA Hellenthal, RL Palma, et al.  \"The chewing lice: World checklist and biological overview. Illinois Natural History Survey Special Publication 24.\"  501 (2003)"},{"key":"22","doi-asserted-by":"crossref","unstructured":"TD Galloway  and RJ Lamb  \"Annu Rev Entomol.\"  66, \"Population dynamics of chewing lice (Phthiraptera) infesting birds (Aves).\" 209 (2021)","DOI":"10.1146\/annurev-ento-041420-075608"},{"key":"42","doi-asserted-by":"crossref","unstructured":"J Potti  and S Merino  \"J Avian Biol.\"  26, \"Louse loads of pied flycatchers: Effects of host's sex, age, condition and relatedness.\" 203 (1995)","DOI":"10.2307\/3677320"}],"container-title":["Journal of Wildlife Diseases"],"original-title":[],"deposited":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T21:03:09Z","timestamp":1791234189000},"score":1,"resource":{"primary":{"URL":"https:\/\/bioone.org\/journals\/journal-of-wildlife-diseases\/volume-62\/issue-2\/JWD-D-25-00151\/PREVALENCE-AND-INTENSITY-OF-PARASITIC-INSECTS-ON-PUERTO-RICAN-BIRDS\/10.7589\/JWD-D-25-00151.full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,20]]},"references-count":62,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2026,4,1]]}},"URL":"https:\/\/doi.org\/10.7589\/jwd-d-25-00151","relation":{},"ISSN":["0090-3558","1943-3700"],"issn-type":[{"value":"0090-3558","type":"print"},{"value":"1943-3700","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,20]]}}}