{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T23:23:10Z","timestamp":1787613790914,"version":"build-2736575974"},"reference-count":69,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T00:00:00Z","timestamp":1781136000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T00:00:00Z","timestamp":1787616000000},"content-version":"vor","delay-in-days":75,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>The crocodile monitor is an arboreal predator, and its ocular adnexa remain poorly characterized. This study provides the first detailed histochemical characterization of the ocular adnexa in this species, including the upper and lower eyelids, the nictitating membrane, the Harderian gland, and the lacrimal gland, based on two adult individuals. Because the available material was limited, the findings should be interpreted as descriptive baseline observations. Histological slides were analyzed using microanatomical assessment, histochemistry, and Picrosirius Red staining under polarized light. Both eyelids were covered by scales and contained a pigmented corneous \u03b2-protein\u2013rich epidermis. A collagen-rich dermis with a thick stratum compactum and an osteoderm located between the dermis and hypodermis were identified. The posterior palpebral epithelium was non-keratinized and contained abundant goblet cells producing mainly acidic mucins. The lower eyelid also contained a dense regular tarsal plate and lymphoid tissue. The nictitating membrane was folded and pigmented, contained hyaline cartilage, and showed conjunctiva-associated lymphoid tissue. Birefringence patterns suggested Type I\u2013like collagen in the perichondrium and Type III\u2013like collagen in the cartilage matrix. The Harderian gland, a large acinar gland, opened onto the bulbar surface of the nictitating membrane and secreted mainly acidic mucins with a moderate neutral mucin component. The small lacrimal gland produced predominantly acidic mucins without detectable neutral mucins. These findings provide a baseline for the protective and secretory ocular structures of the crocodile monitor and represent the first detailed histochemical characterization of the studied orbital glands. In addition, this study contributes to comparative anatomy, veterinary medicine, and exotic animal conservation.<\/jats:p>","DOI":"10.1038\/s41598-026-57505-2","type":"journal-article","created":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T06:03:11Z","timestamp":1781157791000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Morphological and histochemical characterization of eyelids and orbital glands in the crocodile monitor (Varanus salvadorii)"],"prefix":"10.1038","volume":"16","author":[{"given":"Joanna","family":"Kle\u0107kowska-Nawrot","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aleksander","family":"Chr\u00f3szcz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shaw","family":"Badenhorst","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Konrad","family":"Cyprych","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Abit","family":"Akta\u015f","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wojciech","family":"Paszta","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Karolina","family":"Go\u017adziewska-Har\u0142ajczuk","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dominik","family":"Poradowski","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,6,11]]},"reference":[{"key":"57505_CR1","doi-asserted-by":"crossref","unstructured":"Pianka, E. R., King, D. R. & King, R. A. Varanoid Lizards of the World. (Indiana University Press, Bloomington & Indianapolis, 2004).","DOI":"10.2307\/j.ctt2005wjp"},{"key":"57505_CR2","doi-asserted-by":"crossref","unstructured":"Mertens, R. Liste der rezenten Amphibien und Reptilien. Helodermatidae, Varanidae, Lanthanotidae. Das Tierreich 79, i-x, 1\u201326 (1963).","DOI":"10.1515\/9783111385556"},{"key":"57505_CR3","doi-asserted-by":"publisher","first-page":"853","DOI":"10.1098\/rsbl.2012.0460","volume":"8","author":"N Vidal","year":"2012","unstructured":"Vidal, N. et al. Molecular evidence for an Asian origin of monitor lizards followed by Tertiary dispersals to Africa and Australasia. Biol. Lett. 8, 853\u2013855. https:\/\/doi.org\/10.1098\/rsbl.2012.0460 (2012).","journal-title":"Biol. Lett."},{"key":"57505_CR4","doi-asserted-by":"publisher","first-page":"21402","DOI":"10.11609\/jott.7808.14.7.21402-21408","volume":"14","author":"B Reh","year":"2022","unstructured":"Reh, B. & Thomas, J. Range extension and new ecoregion records of the Crocodile Monitor Varanus salvadorii in the Torricelli Mountain Range, Papua New Guinea. J. Threat Taxa. 14, 21402\u201321408. https:\/\/doi.org\/10.11609\/jott.7808.14.7.21402-21408 (2022).","journal-title":"J. Threat Taxa"},{"key":"57505_CR5","unstructured":"Allison, A. Reptiles and Amphibians of the Trans-Fly Region, New Guinea. (Pacific Biological Survey, 2006)."},{"key":"57505_CR6","first-page":"25","volume":"3","author":"HG Horn","year":"2007","unstructured":"Horn, H. G., Sweet, S. S. & Philipp, K. M. On the distribution of the Papuan Monitor (Varanus salvadorii Peters & Doria, 1878) in New Guinea. Adv. Monit. Res. 3, 25\u201343 (2007).","journal-title":"Adv. Monit. Res."},{"key":"57505_CR7","first-page":"4","volume":"1","author":"J Cann","year":"1974","unstructured":"Cann, J. Collecting in Irian Jaya (West New Guinea) during 1972. Bull. Herp Soc. N S W. 1, 4\u201314 (1974).","journal-title":"Bull. Herp Soc. N S W"},{"key":"57505_CR8","unstructured":"Wyneken, J. Reptilian eyes and orbital structures. In Proceedings of the Association of Reptilian and Amphibian Veterinarians 150\u2013162 (2021)."},{"key":"57505_CR9","doi-asserted-by":"crossref","unstructured":"Walls, G. L. The Vertebrate Eye and Its Adaptive Radiation. (Cranbrook Institute of Science, 1942).","DOI":"10.5962\/bhl.title.7369"},{"key":"57505_CR10","doi-asserted-by":"publisher","first-page":"1108","DOI":"10.3390\/ani13061108","volume":"13","author":"T Hellebuyck","year":"2023","unstructured":"Hellebuyck, T. & Vilanova, F. S. Anatomy, physiology, and disorders of the spectacle, subspectacular space, and its lacrimal drainage system in squamates. Animals 13, 1108. https:\/\/doi.org\/10.3390\/ani13061108 (2023).","journal-title":"Animals"},{"key":"57505_CR11","doi-asserted-by":"publisher","unstructured":"Schwab, I. R. & Brooks, D. E. He cries crocodile tears. Br. J. Ophthalmol. 86, 23 https:\/\/doi.org\/10.1136\/bjo.86.1.23 (2002).","DOI":"10.1136\/bjo.86.1.23"},{"key":"57505_CR12","unstructured":"Kardong, K. V. Vertebrates: Comparative Anatomy, Function, and Evolution. 6th edn (McGraw-Hill, 2012)."},{"key":"57505_CR13","doi-asserted-by":"crossref","unstructured":"Montiani-Ferreira, F., Moore, B. A. & Ben-Shlomo, G. Wild and Exotic Animals Ophthalmology. Vol. 1, (Springer, 2022).","DOI":"10.1007\/978-3-030-81273-7"},{"key":"57505_CR14","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1111\/j.1469-7580.1992.tb00378.x","volume":"180","author":"G Chieffi Baccari","year":"1992","unstructured":"Chieffi Baccari, G., Di Matteo, L. & Minucci, S. The orbital glands of the chelonians Pseudemys scripta and Testudo graeca: comparative histological, histochemical and ultrastructural investigations. J. Anat. 180, 1\u201313. https:\/\/doi.org\/10.1111\/j.1469-7580.1992.tb00378.x (1992).","journal-title":"J. Anat."},{"key":"57505_CR15","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1002\/(SICI)1097-0185(199707)248:3<301::AID-AR1>3.0.CO;2-S","volume":"248","author":"SJ Rehorek","year":"1997","unstructured":"Rehorek, S. J. Squamate Harderian gland: an overview. Anat. Rec. 248, 301\u2013306 (1997).","journal-title":"Anat. Rec"},{"key":"57505_CR16","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1002\/(SICI)1097-4687(199703)231:3<253::AID-JMOR4>3.0.CO;2-C","volume":"231","author":"SJ Rehorek","year":"1997","unstructured":"Rehorek, S. J., Firth, B. T. & Hutchinson, M. N. Morphology of the Harderian gland of some Australian geckos. J. Morphol. 231, 253\u2013259 (1997).","journal-title":"J. Morphol."},{"key":"57505_CR17","doi-asserted-by":"publisher","first-page":"706","DOI":"10.1139\/z06-053","volume":"84","author":"SJ Rehorek","year":"2006","unstructured":"Rehorek, S. J., Baker, J. J., Hutchinson, M. N. & Firth, B. T. The Harderian gland of two species of skink (Tiliqua rugosa and Hemiergis decresiensis): significance of lymphatic tissue in the squamate Harderian gland. Can. J. Zool. 84, 706\u2013714. https:\/\/doi.org\/10.1139\/z06-053 (2006).","journal-title":"Can. J. Zool."},{"key":"57505_CR18","unstructured":"Rehorek, S. J. A Comparative Morphological Analysis of Australian Scincid Harderian Glands. Unpublished honours thesis (1992)."},{"key":"57505_CR19","volume-title":"Abstracts of the Second World Congress of Herpetology","author":"SJ Rehorek","year":"1993","unstructured":"Rehorek, S. J., Firth, B. T. & Hutchinson, M. N. A comparative morphological analysis of the Harderian glands of three species of Lerista (Scincidae). In Abstracts of the Second World Congress of Herpetology. (Adelaide, Australia, 1993)."},{"key":"57505_CR20","unstructured":"Stiles, J. & Townsend, W. M. Ophthalmology of Exotic Pets. (Elsevier, 2015)."},{"key":"57505_CR21","doi-asserted-by":"publisher","first-page":"691","DOI":"10.1139\/z71-108","volume":"49","author":"FB Cowan","year":"1971","unstructured":"Cowan, F. B. The ultrastructure of the lachrymal \u2018salt\u2019 gland and the Harderian gland in the euryhaline Malaclemys and some closely related stenohaline emydines. Can. J. Zool. 49, 691\u2013697. https:\/\/doi.org\/10.1139\/z71-108 (1971).","journal-title":"Can. J. Zool."},{"key":"57505_CR22","doi-asserted-by":"crossref","unstructured":"Hillenius, W. J. & Rehorek, S. J. From the eye to the nose: ancient orbital to vomeronasal communication in tetrapods? In Chemical Signals in Vertebrates X (eds Mason, R. T., LeMaster, M. P. & M\u00fcller-Schwarze, D.) 228\u2013241 (Springer, New York, 2005).","DOI":"10.1007\/0-387-25160-X_27"},{"key":"57505_CR23","doi-asserted-by":"publisher","first-page":"1284","DOI":"10.1139\/z00-04","volume":"78","author":"SJ Rehorek","year":"2000","unstructured":"Rehorek, S. J., Hillenius, W. J., Quan, W. & Halpern, M. Passage of the Harderian gland secretions to the vomeronasal organ of the snake, Thamnophis sirtalis. Can. J. Zool. 78, 1284\u20131288. https:\/\/doi.org\/10.1139\/z00-04 (2000).","journal-title":"Can. J. Zool."},{"key":"57505_CR24","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1002\/jmor.10378","volume":"266","author":"SJ Rehorek","year":"2005","unstructured":"Rehorek, S. J., Legenzoff, E. J., Carmody, K., Smith, T. D. & Sedlmayr, J. C. Alligator tears: a re-evaluation of the lacrimal apparatus of the crocodilians. J. Morphol. 266, 298\u2013308. https:\/\/doi.org\/10.1002\/jmor.10378 (2005).","journal-title":"J. Morphol."},{"key":"57505_CR25","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1159\/000305886","volume-title":"Methods and Achievements in Experimental Pathology","author":"SC Spicer","year":"1967","unstructured":"Spicer, S. C. & Henson, J. G. Methods for localizing mucosubstances in epithelial and connective tissue. In Methods and Achievements in Experimental Pathology. (eds (eds Bajusz, E. & Jasmin, F., Karger, S.) 2, 78\u2013112 https:\/\/doi.org\/10.1159\/000305886. (1967)."},{"key":"57505_CR26","unstructured":"International Committee on Veterinary Gross Anatomical Nomenclature. Nomina Anatomica Veterinaria. 6th edn (Editorial Committee, 2017)."},{"key":"57505_CR27","unstructured":"Burck, H. C. Technika Histologiczna. (PZWL, 1975)."},{"key":"57505_CR28","doi-asserted-by":"publisher","first-page":"282","DOI":"10.1016\/j.jsb.2016.03.004","volume":"194","author":"M Calvaresi","year":"2016","unstructured":"Calvaresi, M., Eckhart, L. & Alibardi, L. The molecular organization of the beta-sheet region in corneous beta-proteins (beta-keratins) of sauropsids explains its stability and polymerization into filaments. J. Struct. Biol. 194, 282\u2013291. https:\/\/doi.org\/10.1016\/j.jsb.2016.03.004 (2016).","journal-title":"J. Struct. Biol."},{"key":"57505_CR29","doi-asserted-by":"publisher","first-page":"438","DOI":"10.1002\/jez.b.22840","volume":"330","author":"KB Holthaus","year":"2018","unstructured":"Holthaus, K. B., Eckhart, L., Dalla Valle, L. & Alibardi, L. Review evolution and diversification of corneous beta-proteins, the characteristic epidermal proteins of reptiles and birds. J. Exp. Zool. B Mol. Dev. Evol. 330, 438\u2013453. https:\/\/doi.org\/10.1002\/jez.b.22840 (2018).","journal-title":"J. Exp. Zool. B Mol. Dev. Evol."},{"key":"57505_CR30","doi-asserted-by":"publisher","first-page":"591","DOI":"10.3390\/genes12040591","volume":"12","author":"ADD Parry","year":"2021","unstructured":"Parry, A. D. D. Structures of the \u00df-keratin filaments and keratin intermediate filaments in the epidermal appendages of birds and reptiles (sauropsids). Genes 12, 591. https:\/\/doi.org\/10.3390\/genes12040591 (2021).","journal-title":"Genes"},{"key":"57505_CR31","doi-asserted-by":"publisher","first-page":"751","DOI":"10.1369\/0022155414545787","volume":"62","author":"R Lattouf","year":"2014","unstructured":"Lattouf, R. et al. Picrosirius red staining: a useful tool to appraise collagen networks in normal and pathological tissues. J. Histochem. Cytochem. 62, 751\u2013758. https:\/\/doi.org\/10.1369\/0022155414545787 (2014).","journal-title":"J. Histochem. Cytochem."},{"key":"57505_CR32","doi-asserted-by":"publisher","unstructured":"Rittie, L. (ed) Fibrosis: Methods and Protocols (Methods in Molecular Biology 1627). https:\/\/doi.org\/10.1007\/978-1-4939-7113-8 (Humana Press, 2017).","DOI":"10.1007\/978-1-4939-7113-8"},{"key":"57505_CR33","unstructured":"Cree, A. Tuatara: Biology and Conservation of a Venerable Survivor. (Canterbury University, 2014)."},{"key":"57505_CR34","doi-asserted-by":"publisher","first-page":"3","DOI":"10.3374\/014.053.0101","volume":"53","author":"JA Gauthier","year":"2012","unstructured":"Gauthier, J. A. et al. Assembling the squamate tree of life: perspectives from the phenotype and the fossil record. Bull. Peabody Mus. Nat. Hist. 53, 3\u2013308. https:\/\/doi.org\/10.3374\/014.053.0101 (2012).","journal-title":"Bull. Peabody Mus. Nat. Hist."},{"key":"57505_CR35","doi-asserted-by":"publisher","first-page":"595","DOI":"10.1098\/rstl.1867.0026","volume":"157","author":"A G\u00fcnther","year":"1867","unstructured":"G\u00fcnther, A. Contribution to the anatomy of Hatteria (Rhynchocephalus, Owen). Philos. Trans. R Soc. Lond. 157, 595\u2013629. https:\/\/doi.org\/10.1098\/rstl.1867.0026 (1867).","journal-title":"Philos. Trans. R Soc. Lond."},{"key":"57505_CR36","doi-asserted-by":"crossref","unstructured":"Jacobson, E. R. & Garner, M. M. Infectious Diseases and Pathology of Reptiles: Color Atlas and Text. 2nd edn (CRC, 2020).","DOI":"10.1201\/9780429155567"},{"key":"57505_CR37","doi-asserted-by":"crossref","unstructured":"Underwood, G. Academic Press, London,. The eye. In Biology of the Reptilia Vol. 2: Morphology B (eds Gans, C. & Parsons, T. S.) 1\u201395 (1970).","DOI":"10.2307\/1442337"},{"key":"57505_CR38","doi-asserted-by":"publisher","first-page":"932","DOI":"10.1111\/j.1096-3642.1948.tb00405.x","volume":"118","author":"ADA Bellairs","year":"1948","unstructured":"Bellairs, A. D. A. The eyelids and spectacle in geckos. Proc. Zool. Soc. Lond. 118, 932\u2013942. https:\/\/doi.org\/10.1111\/j.1096-3642.1948.tb00405.x (1948).","journal-title":"Proc. Zool. Soc. Lond."},{"key":"57505_CR39","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1111\/j.1096-3642.1950.tb00949.x","volume":"120","author":"ADA Bellairs","year":"1950","unstructured":"Bellairs, A. D. A. & Boyd, J. D. The lachrymal apparatus in lizards and snakes. II. The anterior part of the lachrymal duct and its relationship with the palate and with the nasal and vomeronasal organs. Proc. Zool. Soc. Lond. 120, 269\u2013310. https:\/\/doi.org\/10.1111\/j.1096-3642.1950.tb00949.x (1950).","journal-title":"Proc. Zool. Soc. Lond."},{"key":"57505_CR40","unstructured":"Robb, J. The Tuatara. (Meadowfield, 1977)."},{"key":"57505_CR41","doi-asserted-by":"publisher","DOI":"10.1016\/B978-0-7020-2782-6.50005-9","volume-title":"Clinical Anatomy and Physiology of Exotic Species","author":"B O\u2019Malley","year":"2005","unstructured":"O\u2019Malley, B. Lizards. In Clinical Anatomy and Physiology of Exotic Species (ed O\u2019Malley, B., Saunders, W. B.) https:\/\/doi.org\/10.1016\/B978-0-7020-2782-6.50005-9. (2005)."},{"key":"57505_CR42","first-page":"533","volume":"33","author":"EM Neher","year":"1935","unstructured":"Neher, E. M. The origin of the brille in Crotalus confluentus lutosus (Great Basin rattlesnake). Trans. Am. Ophthalmol. Soc. 33, 533\u2013545 (1935).","journal-title":"Trans. Am. Ophthalmol. Soc."},{"key":"57505_CR43","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1002\/jmor.20086","volume":"274","author":"L Alibardi","year":"2013","unstructured":"Alibardi, L. Cornification in reptilian epidermis occurs through the deposition of keratin-associated beta-proteins (beta-keratins) onto a scaffold of intermediate filament keratins. J. Morphol. 274, 175\u2013193. https:\/\/doi.org\/10.1002\/jmor.20086 (2013).","journal-title":"J. Morphol."},{"key":"57505_CR44","unstructured":"Ostrowski, K. Histologia. (Wydawnictwo Lekarskie PZWL, 1988)."},{"key":"57505_CR45","doi-asserted-by":"publisher","first-page":"3377","DOI":"10.1021\/pr0702619","volume":"6","author":"M Toni","year":"2007","unstructured":"Toni, M., Dalla Valle, L. & Alibardi, L. Hard (beta-)keratins in the epidermis of reptiles: composition, sequence, and molecular organization. J. Proteome Res. 6, 3377\u20133392. https:\/\/doi.org\/10.1021\/pr0702619 (2007).","journal-title":"J. Proteome Res."},{"key":"57505_CR46","doi-asserted-by":"publisher","first-page":"1352","DOI":"10.1002\/jmor.21004","volume":"280","author":"DC D\u2019Amore","year":"2019","unstructured":"D\u2019Amore, D. C., Moreno, K., McHenry, C. R. & Wroe, S. The role of osteoderms in the evolution and ecology of lizards. J. Morphol. 280, 1352\u20131366. https:\/\/doi.org\/10.1002\/jmor.21004 (2019).","journal-title":"J. Morphol."},{"key":"57505_CR47","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1111\/j.1469-7580.2008.01043.x","volume":"214","author":"MK Vickaryous","year":"2009","unstructured":"Vickaryous, M. K. & Sire, J. Y. The integumentary skeleton of tetrapods: origin, evolution, and development. J. Anat. 214, 441\u2013464. https:\/\/doi.org\/10.1111\/j.1469-7580.2008.01043.x (2009).","journal-title":"J. Anat."},{"key":"57505_CR48","doi-asserted-by":"publisher","first-page":"437","DOI":"10.1002\/aja.1001760406","volume":"176","author":"V Levrat-Calviac","year":"1986","unstructured":"Levrat-Calviac, V. & Zylberberg, L. The structure of the osteoderms in the Gekko: Tarentola mauritanica. Am. J. Anat. 176, 437\u2013446. https:\/\/doi.org\/10.1002\/aja.1001760406 (1986).","journal-title":"Am. J. Anat."},{"key":"57505_CR49","first-page":"1","volume":"185","author":"AP Payne","year":"1994","unstructured":"Payne, A. P. The harderian gland: a tercentennial review. J. Anat. 185, 1\u201349 (1994).","journal-title":"J. Anat."},{"key":"57505_CR50","doi-asserted-by":"publisher","unstructured":"Rios, F. M. & Zimmerman, L. M. Immunology of reptiles. eLS https:\/\/doi.org\/10.1002\/9780470015902.a0025735 (2015).","DOI":"10.1002\/9780470015902.a0025735"},{"key":"57505_CR51","doi-asserted-by":"publisher","first-page":"367","DOI":"10.1111\/azo.12223","volume":"99","author":"J Kle\u0107kowska-Nawrot","year":"2018","unstructured":"Kle\u0107kowska-Nawrot, J. et al. Differences in eyelid microstructure and conjunctiva-associated lymphoid tissue between ornamental and wild birds as a result of habitat adaptation. Acta Zool. 99, 367\u2013394. https:\/\/doi.org\/10.1111\/azo.12223 (2018).","journal-title":"Acta Zool."},{"key":"57505_CR52","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1186\/s12917-025-04677-0","volume":"21","author":"J Kle\u0107kowska-Nawrot","year":"2025","unstructured":"Kle\u0107kowska-Nawrot, J. et al. Descriptive histological analysis of the upper, lower, and third eyelids and the conjunctiva-associated lymphoid tissue in birds of prey. BMC Vet. Res. 21, 213. https:\/\/doi.org\/10.1186\/s12917-025-04677-0 (2025).","journal-title":"BMC Vet. Res."},{"key":"57505_CR53","unstructured":"Schat, K. A., Kaspers, B. & Kaiser, P. Avian Immunology. 2nd edn (Academic, 2014)."},{"key":"57505_CR54","doi-asserted-by":"publisher","first-page":"244","DOI":"10.1016\/j.foodchem.2005.06.053","volume":"99","author":"YK Lin","year":"2006","unstructured":"Lin, Y. K. & Liu, D. C. Comparison of physical\u2013chemical properties of type I collagen from different species. Food Chem. 99, 244\u2013251. https:\/\/doi.org\/10.1016\/j.foodchem.2005.06.053 (2006).","journal-title":"Food Chem."},{"key":"57505_CR55","unstructured":"Pawlina, W. Histology: A Text and Atlas with Correlated Cell and Molecular Biology. (Wolters Kluwer, 2023)."},{"key":"57505_CR56","doi-asserted-by":"publisher","first-page":"18537","DOI":"10.1074\/jbc.M110.112904","volume":"285","author":"JJ Wu","year":"2010","unstructured":"Wu, J. J., Weis, M. A., Kim, L. S. & Eyre, D. R. Type III collagen, a fibril network modifier in articular cartilage. J. Biol. Chem. 285, 18537\u201318544. https:\/\/doi.org\/10.1074\/jbc.M110.112904 (2010).","journal-title":"J. Biol. Chem."},{"key":"57505_CR57","unstructured":"Henning, W. Phylogenetic Systematics (University of Illinois Press, Urbana & Chicago, 1999)."},{"key":"57505_CR58","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1016\/0022-5193(82)90276-4","volume":"97","author":"GV Lauder","year":"1982","unstructured":"Lauder, G. V. Historical biology and the problem of design. J. Theor. Biol. 97, 57\u201367. https:\/\/doi.org\/10.1016\/0022-5193(82)90276-4 (1982).","journal-title":"J. Theor. Biol."},{"key":"57505_CR59","unstructured":"Ridley, M. Evolution. 2nd edn (Oxford University Press, 2004)."},{"key":"57505_CR60","unstructured":"Baumel, J. J., King, A. S., Breazile, J. E., Evans, H. E. & Vanden Berge, J. C. Handbook of Avian Anatomy: Nomina Anatomica Avium 2nd edn (Nuttall Ornithological Club, 1993)."},{"issue":"Suppl. 1","key":"57505_CR61","doi-asserted-by":"publisher","first-page":"30","DOI":"10.1111\/vop.12115","volume":"18","author":"AP Ori\u00e1","year":"2015","unstructured":"Ori\u00e1, A. P. et al. Ophthalmic diagnostic tests, orbital anatomy, and adnexal histology of the broad-snouted caiman (Caiman latirostris). Vet. Ophthalmol. 18 (Suppl. 1), 30\u201339. https:\/\/doi.org\/10.1111\/vop.12115 (2015).","journal-title":"Vet. Ophthalmol."},{"key":"57505_CR62","doi-asserted-by":"publisher","first-page":"1244","DOI":"10.1002\/dvdy.20718","volume":"235","author":"TA Franz-Odendaal","year":"2006","unstructured":"Franz-Odendaal, T. A. & Vickaryous, M. K. Skeletal elements in the vertebrate eye and adnexa\u2014morphological and developmental perspectives. Dev. Dyn. 235, 1244\u20131255. https:\/\/doi.org\/10.1002\/dvdy.20718 (2006).","journal-title":"Dev. Dyn."},{"key":"57505_CR63","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1016\/S1094-9194(01)00007-X","volume":"5","author":"RL Coke","year":"2002","unstructured":"Coke, R. L. & Couillard, N. K. Ocular biology and diseases of Old World chameleons. Vet. Clin. North. Am. Exot Anim. Pract. 5, 275\u2013285. https:\/\/doi.org\/10.1016\/S1094-9194(01)00007-X (2002).","journal-title":"Vet. Clin. North. Am. Exot Anim. Pract."},{"key":"57505_CR64","doi-asserted-by":"publisher","first-page":"299","DOI":"10.1679\/aohc1950.44.299","volume":"44","author":"T Sakai","year":"1981","unstructured":"Sakai, T. The mammalian Harderian gland: morphology, biochemistry, function and phylogeny. Arch. Histol. Jpn. 44, 299\u2013333. https:\/\/doi.org\/10.1679\/aohc1950.44.299 (1981).","journal-title":"Arch. Histol. Jpn"},{"key":"57505_CR65","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1111\/j.1463-5224.2008.00607.x","volume":"11","author":"DK Brudenall","year":"2008","unstructured":"Brudenall, D. K., Schwab, I. R. & Fritsches, K. A. Ocular morphology of the leatherback sea turtle (Dermochelys coriacea). Vet. Ophthalmol. 11, 99\u2013110. https:\/\/doi.org\/10.1111\/j.1463-5224.2008.00607.x (2008).","journal-title":"Vet. Ophthalmol."},{"key":"57505_CR66","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1111\/j.1096-3642.1947.tb00500.x","volume":"117","author":"ADA Bellairs","year":"1947","unstructured":"Bellairs, A. D. A. & Boyd, J. D. The lachrymal apparatus in lizards and snakes. I. The brille, the orbital glands, lachrymal canaliculi and origin of the lachrymal duct. Proc. Zool. Soc. Lond. 117, 81\u2013108. https:\/\/doi.org\/10.1111\/j.1096-3642.1947.tb00500.x (1947).","journal-title":"Proc. Zool. Soc. Lond."},{"key":"57505_CR67","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1002\/ar.1092260302","volume":"226","author":"G Chieffi Baccari","year":"1990","unstructured":"Chieffi Baccari, G., Minucci, S., Di Matteo, L. & Chieffi, G. Harderian gland and the lacrimal gland of the lizard Podarcis s. sicula: histology, histochemistry, and ultrastructure. Anat. Rec. 226, 269\u2013278. https:\/\/doi.org\/10.1002\/ar.1092260302 (1990).","journal-title":"Anat. Rec"},{"key":"57505_CR68","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1002\/(SICI)1097-4687(200005)244:2<137::AID-JMOR4>3.0.CO;2-O","volume":"244","author":"G Chieffi Baccari","year":"2000","unstructured":"Chieffi Baccari, G. et al. Morphology of the Harderian gland of the gecko, Tarentola mauritanica. J. Morphol. 244, 137\u2013142 (2000). 10.1002\/(SICI)1097\u20134687(200005)244:2\u2009<\u2009137:AID-JMOR4\u2009>\u20093.0.CO;2-O","journal-title":"J. Morphol."},{"key":"57505_CR69","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1111\/vop.12184","volume":"18","author":"NM Souza","year":"2015","unstructured":"Souza, N. M. et al. Gross, histologic, and micro-computed tomographic anatomy of the lacrimal system of snakes. Vet. Ophthalmol. 18, 15\u201322. https:\/\/doi.org\/10.1111\/vop.12184 (2015).","journal-title":"Vet. Ophthalmol."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-57505-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-57505-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-57505-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T23:11:21Z","timestamp":1787613081000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-57505-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,11]]},"references-count":69,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["57505"],"URL":"https:\/\/doi.org\/10.1038\/s41598-026-57505-2","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,11]]},"assertion":[{"value":"30 January 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 June 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 June 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"All authors have thoroughly examined and critically evaluated all content of this manuscript, have provided substantial revisions where necessary, and have given their explicit consent for publication in its final form.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"26575"}}