{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T15:49:22Z","timestamp":1788364162683,"version":"build-2803163510"},"reference-count":121,"publisher":"Oxford University Press (OUP)","issue":"6","license":[{"start":{"date-parts":[[2025,5,22]],"date-time":"2025-05-22T00:00:00Z","timestamp":1747872000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/journals\/pages\/open_access\/funder_policies\/chorus\/standard_publication_model"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["2151820"],"award-info":[{"award-number":["2151820"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2025,12,5]]},"abstract":"<jats:title>Synopsis<\/jats:title>\n                  <jats:p>Emerging diseases threaten wildlife worldwide and understanding immune function in the context of the ecology of an organism is critical in predicting disease outcomes. The immune response in reptiles is especially understudied, leaving major gaps in the ability to address disease threats. Freshwater turtles are especially imperiled due to a swath of anthropogenic impacts, including bacterial, viral, and fungal diseases. Because multiple novel pathogen types threaten turtles, understanding nuances in their immune responses may help predict how populations may respond to challenges and shape conservation decisions. We aimed to characterize immune responses following exposure to 3 commercially available antigens representing bacterial, viral, and fungal pathogens in adult red-eared slider turtles (Trachemys scripta elegans) housed in natural conditions. We collected blood samples from red-eared sliders at 0, 12, 24, 48, and 72 h following an antigen treatment (lipopolysaccharide; LPS, polyinosinic-polycytidylic acid; poly(I:C), zymosan, or saline). We created blood smears at 0 and 72 h and the collected serum and buffy layer (serum\u00a0+\u00a0BL) were subjected to 3 manipulations (fresh, frozen, frozen\u00a0+\u00a0heat) for use in microbial killing assays against Escherichia coli, Candida albicans, and Staphylococcus aureus. The combination of antigen treatments on the turtles and manipulations to the collected blood allowed us to examine different immune responses and components. We also quantified corticosterone using enzyme-linked immunosorbent assays to understand energy mobilization following antigen treatment. Microbial killing was generally stable against microbes, regardless of antigen exposure and serum manipulations reveal differences in relative leukocyte and protein-based contribution to killing microbe classes. Overall, the low killing of Gram-positive S. aureus relative to E. coli and C. albicans further suggests that red-eared sliders rely on a robust complement response. Though we did not observe substantial differences among treatments, our results suggest that LPS elicited an increased complement response. The other antigens did not consistently stimulate stronger responses, but more work is needed to understand dose-dependent responses to poly(I:C) and zymosan and in turtles and other reptiles. Overall, our research demonstrates that microbial killing assays can be enhanced through deliberate serum\u00a0+\u00a0BL manipulation and microbe usage as ecoimmunological tools to gain a more robust understanding of the immune response in wild organisms.<\/jats:p>","DOI":"10.1093\/icb\/icaf053","type":"journal-article","created":{"date-parts":[[2025,5,21]],"date-time":"2025-05-21T09:08:20Z","timestamp":1747818500000},"page":"1781-1795","source":"Crossref","is-referenced-by-count":4,"title":["Characterizing Primary Immune Responses Against Three Antigens in a Reptile"],"prefix":"10.1093","volume":"65","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7045-8653","authenticated-orcid":false,"given":"Jennifer","family":"Terry","sequence":"first","affiliation":[{"name":"Environmental Sciences Program, Arkansas State University, State University , Jonesboro, AR 72467 ,","place":["USA"]},{"name":"Arkansas State University, State University Department of Biological Sciences, , Jonesboro, AR 72467 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Isabella V","family":"Davis","sequence":"additional","affiliation":[{"name":"Arkansas State University, State University Department of Biological Sciences, , Jonesboro, AR 72467 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5856-8112","authenticated-orcid":false,"given":"Virginie","family":"Rolland","sequence":"additional","affiliation":[{"name":"Environmental Sciences Program, Arkansas State University, State University , Jonesboro, AR 72467 ,","place":["USA"]},{"name":"Arkansas State University, State University Department of Biological Sciences, , Jonesboro, AR 72467 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0225-5686","authenticated-orcid":false,"given":"Lorin A","family":"Neuman-Lee","sequence":"additional","affiliation":[{"name":"Environmental Sciences Program, Arkansas State University, State University , Jonesboro, AR 72467 ,","place":["USA"]},{"name":"Arkansas State University, State University Department of Biological Sciences, , Jonesboro, AR 72467 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2025,5,22]]},"reference":[{"key":"2025121007115863200_bib1","doi-asserted-by":"crossref","first-page":"e0195617","DOI":"10.1371\/journal.pone.0195617","article-title":"Investigation of multiple mortality events in eastern box turtles (Terrapene carolina carolina)","volume":"13","author":"Adamovicz","year":"2018","journal-title":"PLoS One"},{"key":"2025121007115863200_bib2","first-page":"263","article-title":"Emerging infectious diseases of chelonians: an update","volume":"23","author":"Adamovicz","year":"2020","journal-title":"Vet Clin North Am: Exot Anim Pract"},{"key":"2025121007115863200_bib3","doi-asserted-by":"crossref","first-page":"720","DOI":"10.1002\/jez.2423","article-title":"Plasma complement activation mechanisms differ in ornate (Terrapene ornata ornata) and eastern box turtles (Terrapene carolina carolina)","volume":"333","author":"Adamovicz","year":"2020","journal-title":"J Exp Zool Pt A"},{"key":"2025121007115863200_bib4","doi-asserted-by":"crossref","first-page":"91","DOI":"10.3354\/dao03122","article-title":"Mass mortality of eastern box turtles with upper respiratory disease following atypical cold weather","volume":"124","author":"Agha","year":"2017","journal-title":"Dis Aquat Org"},{"key":"2025121007115863200_bib5","doi-asserted-by":"crossref","first-page":"1192767","DOI":"10.3389\/fimmu.2023.1192767","article-title":"Inhibition of the lectin pathway of complement activation reduces LPS-induced acute respiratory distress syndrome in mice","volume":"14","author":"Ali","year":"2023","journal-title":"Front Immunol"},{"key":"2025121007115863200_bib6","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1111\/j.1365-2435.2010.01759.x","article-title":"The role of constraints and limitations in driving individual variation in immune response","volume":"25","author":"Ardia","year":"2011","journal-title":"Funct Ecol"},{"key":"2025121007115863200_bib7","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1002\/jez.2307","article-title":"Intestinal B cells in the red-eared slider turtle, Trachemys scripta: anatomical distribution and implications for ecological interactions with pathogenic microbes","volume":"331","author":"Ashford","year":"2019","journal-title":"J Exp Zool Pt A"},{"key":"2025121007115863200_bib8","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1002\/jez.2237","article-title":"Antibacterial activities of plasma from the common (Chelydra serpentina) and alligator snapping turtle (Macrochelys temminckii)","volume":"331","author":"Baker","year":"2019","journal-title":"J Exp Zool Pt A"},{"key":"2025121007115863200_bib9","doi-asserted-by":"crossref","first-page":"1","DOI":"10.18637\/jss.v067.i01","article-title":"Fitting linear mixed-effects models using lme4","volume":"67","author":"Bates","year":"2015","journal-title":"J Stat Soft"},{"key":"2025121007115863200_bib10","doi-asserted-by":"crossref","first-page":"2138","DOI":"10.1080\/00949655.2017.1318876","article-title":"Flexible Tweedie regression models for continuous data","volume":"87","author":"Bonat","year":"2017","journal-title":"J Stat Comput Simul"},{"key":"2025121007115863200_bib11","doi-asserted-by":"crossref","first-page":"4930","DOI":"10.4049\/jimmunol.1101790","article-title":"TLR15 is unique to avian and reptilian lineages and recognizes a yeast-derived agonist","volume":"189","author":"Boyd","year":"2012","journal-title":"J Immunol"},{"key":"2025121007115863200_bib12","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/s12979-023-00335-x","article-title":"Sex-specific innate immunity and ageing in long-lived fresh water turtles (Kinosternon flavescens: kinosternidae)","volume":"20","author":"Bronikowski","year":"2023","journal-title":"Immun Ageing"},{"key":"2025121007115863200_bib13","doi-asserted-by":"crossref","first-page":"378","DOI":"10.32614\/RJ-2017-066","article-title":"glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling","volume":"9","author":"Brooks","year":"2017","journal-title":"R J"},{"key":"2025121007115863200_bib15","doi-asserted-by":"crossref","first-page":"cow058","DOI":"10.1093\/conphys\/cow058","article-title":"Parasites, stress and reindeer: infection with abomasal nematodes is not associated with elevated glucocorticoid levels in hair or faeces","volume":"4","author":"Carlsson","year":"2016","journal-title":"Conserv Physiol"},{"key":"2025121007115863200_bib16","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1016\/j.virol.2020.01.016","article-title":"High prevalence of subclinical frog virus 3 infection in freshwater turtles of Ontario, Canada","volume":"543","author":"Carstairs","year":"2020","journal-title":"Virology"},{"key":"2025121007115863200_bib17","doi-asserted-by":"crossref","first-page":"586","DOI":"10.1002\/aqc.3258","article-title":"On a razor\u2019s edge: status and prospects of the critically endangered Bellinger River snapping turtle, Myuchelys georgesi","volume":"30","author":"Chessman","year":"2020","journal-title":"Aquat Conserv"},{"key":"2025121007115863200_bib18","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1093\/icb\/34.3.397","article-title":"Demographics of common snapping turtles (Chelydra serpentina): implications for conservation and management of long-lived organisms","volume":"34","author":"Congdon","year":"1994","journal-title":"Am Zool"},{"key":"2025121007115863200_bib19","doi-asserted-by":"crossref","first-page":"760","DOI":"10.1111\/j.1365-2435.2008.01467.x","article-title":"The use of leukocyte profiles to measure stress in vertebrates: a review for ecologists","volume":"22","author":"Davis","year":"2008","journal-title":"Funct Ecol"},{"key":"2025121007115863200_bib20","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/S0306-4565(00)00026-7","article-title":"Effects of lipopolysaccharide and acclimation temperature on induced behavioral fever in juvenile Iguana iguana","volume":"26","author":"Deen","year":"2001","journal-title":"J Therm Biol"},{"key":"2025121007115863200_bib21","doi-asserted-by":"crossref","first-page":"710","DOI":"10.1111\/j.1365-2656.2011.01813.x","article-title":"Beyond phytohaemagglutinin: assessing vertebrate immune function across ecological contexts","volume":"80","author":"Demas","year":"2011","journal-title":"J Anim Ecol"},{"key":"2025121007115863200_bib22","doi-asserted-by":"crossref","first-page":"1345","DOI":"10.1016\/j.psyneuen.2012.05.008","article-title":"Stress-induced redistribution of immune cells-from barracks to boulevards to battlefields: a tale of three hormones\u2014Curt Richter award winner","volume":"37","author":"Dhabhar","year":"2012","journal-title":"Psychoneuroendocrinology"},{"key":"2025121007115863200_bib24","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1111\/acv.12253","article-title":"Consequences of individual removal on persistence of a protected population of long-lived turtles","volume":"19","author":"Dodd","year":"2016","journal-title":"Anim Conserv"},{"key":"2025121007115863200_bib25","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1007\/s43621-022-00083-w","article-title":"Re-evaluating invasive species in degraded ecosystems: a case study of red-eared slider turtles as partial ecological analogs","volume":"3","author":"Dupuis-Desormeaux","year":"2022","journal-title":"Discov Sustain"},{"key":"2025121007115863200_bib26","doi-asserted-by":"crossref","DOI":"10.56021\/9780801891212","volume-title":"Turtles of the United States and Canada","author":"Ernst","year":"2009","edition":"2nd ed."},{"key":"2025121007115863200_bib27","doi-asserted-by":"crossref","first-page":"7260","DOI":"10.1038\/s41598-024-57792-7","article-title":"A novel multi-variate immunological approach, reveals immune variation associated with environmental conditions, and co-infection in the koala (Phascolarctos cinereus)","volume":"14","author":"Fernandez","year":"2024","journal-title":"Sci Rep"},{"key":"2025121007115863200_bib28","doi-asserted-by":"crossref","first-page":"747","DOI":"10.1590\/S1984-46702009000400020","article-title":"Characterization of innate immune activity in Phrynops geoffroanus (Testudines: chelidae)","volume":"26","author":"Ferronato","year":"2009","journal-title":"Zoologia (Curitiba, Impr"},{"key":"2025121007115863200_bib29","doi-asserted-by":"crossref","first-page":"1557","DOI":"10.1093\/icb\/icac116","article-title":"Reptilian innate immunology and ecoimmunology: what do we know and where are we going?","volume":"62","author":"Field","year":"2022","journal-title":"Integr Comp Biol"},{"key":"2025121007115863200_bib30","doi-asserted-by":"crossref","first-page":"114350","DOI":"10.1016\/j.ygcen.2023.114350","article-title":"Investigating relationships among stress, reproduction, and immunity in three species of watersnake","volume":"343","author":"Field","year":"2023","journal-title":"Gen Comp Endocrinol"},{"key":"2025121007115863200_bib31","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1111\/imr.12472","article-title":"The \u201cins and outs\u201d of complement-driven immune responses","volume":"274","author":"Freeley","year":"2016","journal-title":"Immunol Rev"},{"key":"2025121007115863200_bib32","doi-asserted-by":"crossref","first-page":"482","DOI":"10.1242\/bio.2012919","article-title":"Improved ex vivo method for microbiocidal activity across vertebrate species","volume":"1","author":"French","year":"2012","journal-title":"Biol Open"},{"key":"2025121007115863200_bib33","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/j.molimm.2004.03.026","article-title":"Primitive complement system\u2014recognition and activation","volume":"41","author":"Fujita","year":"2004","journal-title":"Mol Immunol"},{"key":"2025121007115863200_bib34","doi-asserted-by":"crossref","first-page":"111213","DOI":"10.1016\/j.cbpa.2022.111213","article-title":"Immune and endocrine responses of Cururu toads (Rhinella icterica) in their natural habitat after LPS stimulation","volume":"269","author":"Garcia\u00a0Neto","year":"2022","journal-title":"Comp Biochem Physiol A: Mol Integr Physiol"},{"key":"2025121007115863200_bib35","doi-asserted-by":"crossref","first-page":"e12393","DOI":"10.1111\/conl.12393","article-title":"Identifying species conservation strategies to reduce disease-associated declines","volume":"11","author":"Gerber","year":"2018","journal-title":"Conserv Lett"},{"key":"2025121007115863200_bib36","article-title":"Species profile: t rachemys scripta elegans","author":"Global Invasive Species Database","year":"2025"},{"key":"2025121007115863200_bib37","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1086\/688694","article-title":"Seasonal acclimation of constitutive immunity in gopher tortoises Gopherus polyphemus","volume":"89","author":"Goessling","year":"2016","journal-title":"Physiol Biochem Zool"},{"key":"2025121007115863200_bib38","doi-asserted-by":"crossref","first-page":"484","DOI":"10.1086\/692116","article-title":"More than fever: thermoregulatory responses to immunological stimulation and consequences of thermoregulatory strategy on innate immunity in gopher tortoises (Gopherus polyphemus )","volume":"90","author":"Goessling","year":"2017","journal-title":"Physiol Biochem Zool"},{"key":"2025121007115863200_bib39","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1002\/jez.2252","article-title":"Rapid thermal immune acclimation in common musk turtles (Sternotherus odoratus)","volume":"331","author":"Goessling","year":"2019","journal-title":"J Exp Zool Pt A"},{"key":"2025121007115863200_bib40","doi-asserted-by":"crossref","first-page":"113269","DOI":"10.1016\/j.ygcen.2019.113269","article-title":"Relationships between glucocorticoids and infection with Batrachochytrium dendrobatidis in three amphibian species","volume":"285","author":"Hammond","year":"2020","journal-title":"Gen Comp Endocrinol"},{"key":"2025121007115863200_bib41","doi-asserted-by":"crossref","first-page":"1389","DOI":"10.1002\/joc.2162","article-title":"Two Tweedie distributions that are near-optimal for modelling monthly rainfall in Australia","volume":"31","author":"Hasan","year":"2011","journal-title":"Intl J Climatol"},{"key":"2025121007115863200_bib42","doi-asserted-by":"crossref","first-page":"780","DOI":"10.1111\/j.1095-8649.2007.01730.x","article-title":"Cortisol responses of pallid sturgeon and yellow perch following challenge with lipopolysaccharide","volume":"72","author":"Haukenes","year":"2008","journal-title":"J Fish Biol"},{"key":"2025121007115863200_bib43","doi-asserted-by":"crossref","first-page":"75","DOI":"10.3354\/dao02812","article-title":"First reported outbreak of severe spirorchiidiasis in Emys orbicularis, probably resulting from a parasite spillover event","volume":"113","author":"Iglesias","year":"2015","journal-title":"Dis Aquat Org"},{"key":"2025121007115863200_bib44","doi-asserted-by":"crossref","first-page":"632","DOI":"10.1086\/422397","article-title":"Role of mannose-binding lectin in the innate defense against Candida albicans: enhancement of complement activation, but lack of opsonic function, in phagocytosis","volume":"190","author":"Ip","year":"2004","journal-title":"J Infect Dis"},{"key":"2025121007115863200_bib45","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1577\/1548-8667(2003)015<0001:CEONAA>2.0.CO;2","article-title":"Cumulative effects of natural and anthropogenic stress on immune function and disease resistance in juvenile chinook salmon","volume":"15","author":"Jacobson","year":"2003","journal-title":"J Aquat Animal Health"},{"key":"2025121007115863200_bib46","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1146\/annurev.immunol.20.083001.084359","article-title":"Innate immune recognition","volume":"20","author":"Janeway","year":"2002","journal-title":"Annu Rev Immunol"},{"key":"2025121007115863200_bib47","doi-asserted-by":"crossref","first-page":"958","DOI":"10.1643\/h2020151","article-title":"Turtle shell disease fungus (Emydomyces testavorans): first documented occurrence in California and prevalence in free-living turtles","volume":"109","author":"Lambert","year":"2021","journal-title":"Ichthyol Herpetol"},{"key":"2025121007115863200_bib48","doi-asserted-by":"crossref","first-page":"132","DOI":"10.1038\/nrmicro1824","article-title":"Complement evasion by human pathogens","volume":"6","author":"Lambris","year":"2008","journal-title":"Nat Rev Micro"},{"key":"2025121007115863200_bib49","article-title":"emmeans: estimated marginal means, aka least-squares means","author":"Lenth","year":"2022"},{"key":"2025121007115863200_bib50","doi-asserted-by":"crossref","first-page":"jeb225185","DOI":"10.1242\/jeb.225185","article-title":"The metabolic response to an immune challenge in a viviparous snake,\u00a0Sistrurus miliarius","volume":"223","author":"Lind","year":"2020","journal-title":"J Exp Biol"},{"key":"2025121007115863200_bib51","doi-asserted-by":"crossref","first-page":"20150457","DOI":"10.1098\/rstb.2015.0457","article-title":"Snake fungal disease: an emerging threat to wild snakes","volume":"371","author":"Lorch","year":"2016","journal-title":"Philos Trans R Soc B"},{"key":"2025121007115863200_bib52","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1016\/j.ygcen.2016.06.008","article-title":"Changes in corticosterone concentrations and behavior during Mycoplasma gallisepticum infection in house finches (Haemorhous mexicanus)","volume":"235","author":"Love","year":"2016","journal-title":"Gen Comp Endocrinol"},{"key":"2025121007115863200_bib53","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1093\/biosci\/biy095","article-title":"Where have all the turtles gone, and why does it matter?","volume":"68","author":"Lovich","year":"2018","journal-title":"BioSci"},{"key":"2025121007115863200_bib54","doi-asserted-by":"crossref","first-page":"2579","DOI":"10.3389\/fimmu.2018.02579","article-title":"The influence of parasite infections on host immunity to co-infection with other pathogens","volume":"9","author":"Mabbot","year":"2018","journal-title":"Front Immunol"},{"key":"2025121007115863200_bib55","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1056\/NEJMra2206346","article-title":"Humoral innate immunity and acute-phase proteins","volume":"388","author":"Mantovani","year":"2023","journal-title":"N Engl J Med"},{"key":"2025121007115863200_bib56","doi-asserted-by":"crossref","first-page":"1234","DOI":"10.3382\/ps.2011-01389","article-title":"Body temperature responses of Pekin ducks (Anas platyrhynchos domesticus) exposed to different pathogens","volume":"90","author":"Marais","year":"2011","journal-title":"Poult Sci"},{"key":"2025121007115863200_bib57","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1098\/rstb.2007.2142","article-title":"Seasonal changes in vertebrate immune activity: mediation by physiological trade-offs","volume":"363","author":"Martin","year":"2008","journal-title":"Philos Trans R Soc B"},{"key":"2025121007115863200_bib58","doi-asserted-by":"crossref","first-page":"e0230760","DOI":"10.1371\/journal.pone.0230760","article-title":"Disease risk analysis in sea turtles: a baseline study to inform conservation efforts","volume":"15","author":"Mashkour","year":"2020","journal-title":"PLoS One"},{"key":"2025121007115863200_bib59","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3389\/fevo.2017.00028","article-title":"Success of wildlife disease treatment depends on host immune response","volume":"5","author":"Maslo","year":"2017","journal-title":"Front Ecol Evol"},{"key":"2025121007115863200_bib60","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1016\/j.dci.2004.07.006","article-title":"A hemolysis-hemagglutination assay for characterizing constitutive innate humoral immunity in wild and domestic birds","volume":"29","author":"Matson","year":"2005","journal-title":"Dev Compar Immunol"},{"key":"2025121007115863200_bib61","doi-asserted-by":"crossref","first-page":"805","DOI":"10.1016\/j.addr.2007.11.005","article-title":"TLR3: interferon induction by double-stranded RNA including poly(I:C)","volume":"60","author":"Matsumoto","year":"2008","journal-title":"Adv Drug Deliv Rev"},{"key":"2025121007115863200_bib62","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/j.ygcen.2015.07.008","article-title":"How do duration, frequency, and intensity of exogenous CORT elevation affect immune outcomes of stress?","volume":"222","author":"McCormick","year":"2015","journal-title":"Gen Comp Endocrinol"},{"key":"2025121007115863200_bib63","doi-asserted-by":"crossref","first-page":"488","DOI":"10.1016\/j.cbpa.2006.01.009","article-title":"Characterization of serum complement activity of saltwater (Crocodylus porosus) and freshwater (Crocodylus johnstoni) crocodiles","volume":"143","author":"Merchant","year":"2006","journal-title":"Comp Biochem Physiol A: Mol Integr Physiol"},{"key":"2025121007115863200_bib64","first-page":"300","article-title":"Differential protein expression in alligator leukocytes in response to bacterial lipopolysaccharide injection","volume":"4","author":"Merchant","year":"2009","journal-title":"Comp Biochem Physiol D: Genom Proteom"},{"key":"2025121007115863200_bib65","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1016\/j.dci.2006.05.013","article-title":"Quantifying and comparing constitutive immunity across avian species","volume":"31","author":"Millet","year":"2007","journal-title":"Dev Compar Immunol"},{"key":"2025121007115863200_bib66","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1638\/2013-0147R1.1","article-title":"Effects of ranavirus infection of red-eared sliders (Trachemys scripta elegans) on plasma proteins","volume":"45","author":"Moore","year":"2014","journal-title":"J Zoo Wildl Med"},{"key":"2025121007115863200_bib67","doi-asserted-by":"crossref","first-page":"108496","DOI":"10.1016\/j.biocon.2020.108496","article-title":"Predation and disease limit population recovery following 15 years of headstarting an endangered freshwater turtle","volume":"245","author":"Mullin","year":"2020","journal-title":"Biol Conserv"},{"key":"2025121007115863200_bib68","doi-asserted-by":"crossref","first-page":"178","DOI":"10.1643\/CP-02-058R2","article-title":"Seasonal changes in lymphoid distribution of the turtle Mauremys caspica","volume":"2004","author":"Mu\u00f1oz","year":"2004","journal-title":"Copeia"},{"key":"2025121007115863200_bib69","first-page":"327","article-title":"A simple and reliable system for marking hard-shelled turtles: the North American code","volume":"48","author":"Nagle","year":"2017","journal-title":"Herpetol Rev"},{"key":"2025121007115863200_bib70","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1086\/700396","article-title":"Interrelations among multiple metrics of immune and physiological function in a squamate, the common gartersnake (Thamnophis sirtalis)","volume":"92","author":"Neuman-Lee","year":"2019","journal-title":"Physiol Biochem Zool"},{"key":"2025121007115863200_bib71","doi-asserted-by":"crossref","first-page":"coab074","DOI":"10.1093\/conphys\/coab074","article-title":"Applied ecoimmunology: using immunological tools to improve conservation efforts in a changing world","volume":"9","author":"Ohmer","year":"2021","journal-title":"Conserv Physiol"},{"key":"2025121007115863200_bib72","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1002\/jez.2137","article-title":"Immune variation during pregnancy suggests immune component-specific costs of reproduction in a viviparous snake with disparate life-history strategies","volume":"327","author":"Palacios","year":"2017","journal-title":"J Exp Zool Pt A"},{"key":"2025121007115863200_bib73","doi-asserted-by":"crossref","first-page":"5897","DOI":"10.4049\/jimmunol.170.12.5897","article-title":"B cells activated by lipopolysaccharide, but not by anti-Ig and anti-CD40 antibody, induce anergy in CD8 T Cells: role of TGF-1 1 B cells","volume":"170","author":"Parekh","year":"2003","journal-title":"J Immunol"},{"key":"2025121007115863200_bib74","doi-asserted-by":"crossref","first-page":"e66","DOI":"10.1038\/emm.2013.97","article-title":"Recognition of lipopolysaccharide pattern by TLR4 complexes","volume":"45","author":"Park","year":"2013","journal-title":"Exp Mol Med"},{"key":"2025121007115863200_bib75","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1111\/j.1600-0706.2011.19735.x","article-title":"Population-level influence of a recurring disease on a long-lived wildlife host","volume":"121","author":"Perez-Heydrich","year":"2012","journal-title":"Oikos"},{"key":"2025121007115863200_bib76","doi-asserted-by":"crossref","first-page":"861","DOI":"10.3390\/ani11030861","article-title":"Insights on immune function in free-ranging green sea turtles (Chelonia mydas) with and without fibropapillomatosis","volume":"11","author":"Perrault","year":"2021","journal-title":"Animals"},{"key":"2025121007115863200_bib77","doi-asserted-by":"crossref","first-page":"20976","DOI":"10.1038\/s41598-020-77865-7","article-title":"Cross-continental emergence of Nannizziopsis barbatae disease may threaten wild Australian lizards","volume":"10","author":"Peterson","year":"2020","journal-title":"Sci Rep"},{"key":"2025121007115863200_bib78","doi-asserted-by":"crossref","first-page":"916","DOI":"10.1016\/S0306-4530(02)00108-7","article-title":"Two formulas for computation of the area under the curve represent measures of total hormone concentration versus time-dependent change","volume":"28","author":"Pruessner","year":"2003","journal-title":"Psychoneuroendocrinology"},{"key":"2025121007115863200_bib79","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/j.dci.2019.05.005","article-title":"Diversity, immunoregulatory action and structure-activity relationship of green sea turtle cathelicidins","volume":"98","author":"Qiao","year":"2019","journal-title":"Dev Compar Immunol"},{"key":"2025121007115863200_bib80","volume-title":"R: a language and environment for statistical computing","author":"R Core Team","year":"2022"},{"key":"2025121007115863200_bib81","doi-asserted-by":"crossref","first-page":"7853","DOI":"10.1074\/jbc.M707591200","article-title":"Activation of complement component C5: comparison of C5 convertases of the lectin pathway and the classical pathway of complement","volume":"283","author":"Rawal","year":"2008","journal-title":"J Biol Chem"},{"key":"2025121007115863200_bib82","doi-asserted-by":"crossref","first-page":"e0241414","DOI":"10.1371\/journal.pone.0241414","article-title":"Temperature affects the host hematological and cytokine response following experimental ranavirus infection in red-eared sliders (Trachemys scripta elegans)","volume":"15","author":"Rayl","year":"2020","journal-title":"PLoS One"},{"key":"2025121007115863200_bib83","first-page":"1","article-title":"Turtles of the world: annotated checklist and atlas of taxonomy, synonymy, distribution, and conservation status","volume-title":"Conservation biology of freshwater turtles and tortoises: a compilation project of the IUCN\/SSC tortoise and freshwater turtle specialist group","author":"Turtle Taxonomy Working Group","year":"2021","edition":"9th ed"},{"key":"2025121007115863200_bib84","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1189\/jlb.68.6.785","article-title":"Leukocyte antimicrobial peptides: multifunctional effector molecules of innate immunity","volume":"68","author":"Risso","year":"2000","journal-title":"J Leukoct Biol"},{"key":"2025121007115863200_bib85","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.cbpb.2004.11.004","article-title":"Collecting baseline corticosterone samples in the field: is under 3 min good enough?","volume":"140","author":"Romero","year":"2005","journal-title":"Comp Biochem Physiol A: Mol Integr Physiol"},{"key":"2025121007115863200_bib86","doi-asserted-by":"crossref","first-page":"105322","DOI":"10.1016\/j.dci.2025.105322","article-title":"Memory responses to ovalbumin-immunization to Mojave desert tortoises","volume":"164","author":"Sandmeier","year":"2025","journal-title":"Dev Compar Immunol"},{"key":"2025121007115863200_bib87","doi-asserted-by":"crossref","first-page":"1255","DOI":"10.1016\/j.biocon.2009.02.001","article-title":"Upper respiratory tract disease (URTD) as a threat to desert tortoise populations: a reevaluation","volume":"142","author":"Sandmeier","year":"2009","journal-title":"Biol Conserv"},{"key":"2025121007115863200_bib88","doi-asserted-by":"crossref","first-page":"1078","DOI":"10.1242\/bio.20122527","article-title":"A trade-off between natural and acquired antibody production in a reptile: implications for long-term resistance to disease","volume":"1","author":"Sandmeier","year":"2012","journal-title":"Biol Open"},{"key":"2025121007115863200_bib89","first-page":"55","article-title":"How do glucocorticoids influence stress responses? Integrating permissive, suppressive, stimulatory, and preparative actions","volume":"21","author":"Sapolsky","year":"2000","journal-title":"Endocr Rev"},{"key":"2025121007115863200_bib90","doi-asserted-by":"crossref","first-page":"417","DOI":"10.4049\/jimmunol.171.1.417","article-title":"Direct binding of toll-like receptor 2 to zymosan, and zymosan-induced NF-B activation and TNF-secretion are down-regulated by lung collectin surfactant protein A","volume":"171","author":"Sato","year":"2003","journal-title":"J Immunol"},{"key":"2025121007115863200_bib91","doi-asserted-by":"crossref","first-page":"e524","DOI":"10.1111\/csp2.524","article-title":"Conservation translocations for amphibian species threatened by chytrid fungus: a review, conceptual framework, and recommendations","volume":"3","author":"Scheele","year":"2021","journal-title":"Conserv Sci Pract"},{"key":"2025121007115863200_bib92","doi-asserted-by":"crossref","first-page":"1420","DOI":"10.1016\/j.biocon.2009.02.007","article-title":"Magnitude of the US trade in amphibians and presence of Batrachochytrium dendrobatidis and ranavirus infection in imported North American bullfrogs (Rana catesbeiana)","volume":"142","author":"Schloegel","year":"2009","journal-title":"Biol Conserv"},{"key":"2025121007115863200_bib93","doi-asserted-by":"crossref","first-page":"3095","DOI":"10.1007\/s10531-020-02021-6","article-title":"Biodiversity loss, emerging pathogens, and human health risks","volume":"29","author":"Schmeller","year":"2020","journal-title":"Biodivers Conserv"},{"key":"2025121007115863200_bib94","doi-asserted-by":"crossref","first-page":"15259","DOI":"10.1038\/s41598-022-18240-6","article-title":"Differences in acute phase response to bacterial, fungal and viral antigens in greater mouse-eared bats (Myotis myotis)","volume":"12","author":"Seltmann","year":"2022","journal-title":"Sci Rep"},{"key":"2025121007115863200_bib95","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1016\/j.dci.2018.11.010","article-title":"Roles of polymorphic cathelicidins in innate immunity of soft-shell turtle, Pelodiscus sinensis","volume":"92","author":"Shi","year":"2019","journal-title":"Dev Compar Immunol"},{"key":"2025121007115863200_bib96","doi-asserted-by":"crossref","first-page":"37","DOI":"10.3354\/dao02663","article-title":"Prevalence of selected pathogens in western pond turtles and sympatric introduced red-eared sliders in California, USA","volume":"107","author":"Silbernagel","year":"2013","journal-title":"Dis Aquat Org"},{"key":"2025121007115863200_bib97","doi-asserted-by":"crossref","first-page":"1279","DOI":"10.1128\/iai.38.3.1279-1284.1982","article-title":"Inhibition of zymosan-induced alternative complement pathway activation by concanavalin A","volume":"38","author":"Smith","year":"1982","journal-title":"Infect Immun"},{"key":"2025121007115863200_bib98","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1016\/j.biocon.2018.03.009","article-title":"Profiling a possible rapid extinction event in a long-lived species","volume":"221","author":"Spencer","year":"2018","journal-title":"Biol Conserv"},{"key":"2025121007115863200_bib99","doi-asserted-by":"crossref","first-page":"761","DOI":"10.7589\/JWD-D-20-00204","article-title":"Evaluation of immune function in two populations of green sea turtles (Chelonia mydas) in a degraded versus a nondegraded habitat","volume":"57","author":"Sposato","year":"2021","journal-title":"J Wildl Dis"},{"key":"2025121007115863200_bib100","first-page":"944","article-title":"History of predator exposure affects cell-mediated immunity in female eastern fence lizards, Sceloporus undulatus (Squamata: phrynosomatidae)","volume":"128","author":"Sprayberry","year":"2019","journal-title":"Biol J Linn Soc"},{"key":"2025121007115863200_bib101","doi-asserted-by":"crossref","first-page":"R721","DOI":"10.1016\/j.cub.2020.04.088","article-title":"Turtles and tortoises are in trouble","volume":"30","author":"Stanford","year":"2020","journal-title":"Curr Biol"},{"key":"2025121007115863200_bib102","first-page":"497","article-title":"Opsonization of various capsular (K) E. coli by the alternative complement pathway","volume":"50","author":"Stevens","year":"1983","journal-title":"Immunol"},{"key":"2025121007115863200_bib103","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1016\/B978-0-7020-4089-4.00023-8","article-title":"Immunological principles: antigens and antigen recognition","volume-title":"Medical microbiology","author":"Stewart","year":"2012","edition":"18th ed"},{"key":"2025121007115863200_bib104","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1002\/jez.2111","article-title":"Relationships between parasitic infection and natural antibodies, age, and sex in a long-lived vertebrate","volume":"327","author":"Stromsland","year":"2017","journal-title":"J Exp Zool Pt A"},{"key":"2025121007115863200_bib105","doi-asserted-by":"crossref","first-page":"396","DOI":"10.1016\/j.micres.2013.02.005","article-title":"Innate immune recognition of microbial cell wall components and microbial strategies to evade such recognitions","volume":"168","author":"Sukhithasri","year":"2013","journal-title":"Microbiol Res"},{"key":"2025121007115863200_bib106","doi-asserted-by":"crossref","first-page":"114288","DOI":"10.1016\/j.ygcen.2023.114288","article-title":"Assessment of glucocorticoids, sex steroids, and innate immunity in red-eared slider turtles (Trachemys scripta elegans)","volume":"339","author":"Terry","year":"2023","journal-title":"Gen Comp Endocrinol"},{"key":"2025121007115863200_bib107","doi-asserted-by":"crossref","first-page":"e0126155","DOI":"10.1371\/journal.pone.0126155","article-title":"Oxidant trade-offs in immunity: an experimental test in a lizard","volume":"10","author":"Tobler","year":"2015","journal-title":"PLoS One"},{"key":"2025121007115863200_bib108","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1016\/j.pt.2015.01.007","article-title":"Emerging infectious diseases of wildlife: a critical perspective","volume":"31","author":"Tompkins","year":"2015","journal-title":"Trends Parasitol"},{"key":"2025121007115863200_bib109","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1093\/beheco\/ari091","article-title":"Age, parasites, and condition affect humoral immune response in tropical pythons","volume":"17","author":"Ujvari","year":"2006","journal-title":"Behav Ecol"},{"key":"2025121007115863200_bib110","doi-asserted-by":"crossref","first-page":"176","DOI":"10.1177\/09680519030090030601","article-title":"Macrophage recognition of zymosan particles","volume":"9","author":"Underhill","year":"2003","journal-title":"J Endotoxin Res"},{"key":"2025121007115863200_bib111","doi-asserted-by":"crossref","first-page":"723","DOI":"10.3390\/ph7060723","article-title":"Antimicrobial peptides in reptiles","volume":"7","author":"Van\u00a0Hoek","year":"2014","journal-title":"Pharmaceuticals"},{"key":"2025121007115863200_bib112","doi-asserted-by":"crossref","first-page":"jeb24458","DOI":"10.1242\/jeb.244583","article-title":"The acute phase response in bats (Carollia perspicillata) varies with time and dose of the immune challenge","volume":"225","author":"Viola","year":"2022","journal-title":"J Exp Biol"},{"key":"2025121007115863200_bib113","doi-asserted-by":"crossref","first-page":"2751","DOI":"10.4049\/jimmunol.0903201","article-title":"Cutting edge: polyinosinic:polycytidylic acid boosts the generation of memory CD8 T Cells through melanoma differentiation-associated protein 5 expressed in stromal cells","volume":"184","author":"Wang","year":"2010","journal-title":"J Immunol"},{"key":"2025121007115863200_bib114","doi-asserted-by":"crossref","first-page":"346","DOI":"10.1038\/86354","article-title":"Leptospiral lipopolysaccharide activates cells through a TLR2-dependent mechanism","volume":"2","author":"Werts","year":"2001","journal-title":"Nat Immunol"},{"key":"2025121007115863200_bib116","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1644\/BHE-004.1","article-title":"The concept of allostasis: coping with a capricious environment","volume":"86","author":"Wingfield","year":"2005","journal-title":"J Mammal"},{"key":"2025121007115863200_bib117","doi-asserted-by":"crossref","first-page":"578","DOI":"10.1177\/0300985820985217","article-title":"Shell lesions associated with Emydomyces testavorans infection in freshwater aquatic turtles","volume":"58","author":"Woodburn","year":"2021","journal-title":"Vet Pathol"},{"key":"2025121007115863200_bib118","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/j.cellimm.2010.03.020","article-title":"Cytokine release of human NK cells solely triggered with polyI:C","volume":"263","author":"Wulff","year":"2010","journal-title":"Cell Immunol"},{"key":"2025121007115863200_bib119","doi-asserted-by":"crossref","first-page":"1572","DOI":"10.1093\/icb\/icac022","article-title":"Adaptive immunity in reptiles: conventional components but unconventional strategies","volume":"62","author":"Zimmerman","year":"2022","journal-title":"Integr Comp Biol"},{"key":"2025121007115863200_bib120","doi-asserted-by":"crossref","first-page":"1061","DOI":"10.1111\/1365-2435.12273","article-title":"A vertebrate cytokine primer for eco-immunologists","volume":"28","author":"Zimmerman","year":"2014","journal-title":"Funct Ecol"},{"key":"2025121007115863200_bib121","doi-asserted-by":"crossref","first-page":"1383","DOI":"10.1111\/1365-2435.12867","article-title":"Immunocompetence in a long-lived ectothermic vertebrate is temperature dependent but shows no decline in older adults","volume":"31","author":"Zimmerman","year":"2017","journal-title":"Funct Ecol"},{"key":"2025121007115863200_bib122","first-page":"633","article-title":"Humoral immune responses are maintained with age in a long-lived ectotherm, the red-eared slider turtle","volume":"216","author":"Zimmerman","year":"2013","journal-title":"J Exp Biol"},{"key":"2025121007115863200_bib123","doi-asserted-by":"crossref","first-page":"1477","DOI":"10.1242\/jeb.037770","article-title":"Variation in the seasonal patterns of innate and adaptive immunity in the red-eared slider (Trachemys scripta)","volume":"213","author":"Zimmerman","year":"2010","journal-title":"J Exp Biol"},{"key":"2025121007115863200_bib124","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1242\/jeb.038315","article-title":"Understanding the vertebrate immune system: insights from the reptilian perspective","volume":"213","author":"Zimmerman","year":"2010","journal-title":"J Exp Biol"}],"container-title":["Integrative And Comparative Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/icb\/advance-article-pdf\/doi\/10.1093\/icb\/icaf053\/63296313\/icaf053.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/icb\/article-pdf\/65\/6\/1781\/63296313\/icaf053.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/icb\/article-pdf\/65\/6\/1781\/63296313\/icaf053.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,10]],"date-time":"2025-12-10T12:12:24Z","timestamp":1765368744000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/icb\/article\/65\/6\/1781\/8140839"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,5,22]]},"references-count":121,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2025,5,22]]},"published-print":{"date-parts":[[2025,12,5]]}},"URL":"https:\/\/doi.org\/10.1093\/icb\/icaf053","relation":{},"ISSN":["1540-7063","1557-7023"],"issn-type":[{"value":"1540-7063","type":"print"},{"value":"1557-7023","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2025,12]]},"published":{"date-parts":[[2025,5,22]]}}}