{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,8]],"date-time":"2026-08-08T14:45:07Z","timestamp":1786200307535,"version":"build-2736575974"},"reference-count":65,"publisher":"MDPI AG","issue":"11","license":[{"start":{"date-parts":[[2023,11,17]],"date-time":"2023-11-17T00:00:00Z","timestamp":1700179200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Dreamworld Wildlife Foundation donation and the Annandale Bequest, School of Veterinary Science, The University of Queensland"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biology"],"abstract":"<jats:p>The koala (Phascolarctos cinereus), while being an iconic Australian marsupial, has recently been listed as endangered. To establish an improved understanding of normal reproductive anatomy, this paper brings together unpublished research which has approached the topic from two perspectives: (1) the establishment of an artificial insemination program, and (2) the definition of Chlamydia spp.-derived histopathological changes of the female koala urogenital system. Based on the presentation and histological processing of over 70 opportunistic specimens, recovered from wildlife hospitals in Southeast Queensland (Australia), we describe the gross and microanatomy of the koala ovary, oviduct, uteri, vaginal complex, and urogenital sinus during the interestrous, proliferative, and luteal phases of the reproductive cycle.<\/jats:p>","DOI":"10.3390\/biology12111445","type":"journal-article","created":{"date-parts":[[2023,11,17]],"date-time":"2023-11-17T09:10:38Z","timestamp":1700212238000},"page":"1445","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":10,"title":["Anatomy of the Female Koala Reproductive Tract"],"prefix":"10.3390","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1415-0299","authenticated-orcid":false,"given":"Sara","family":"Pagliarani","sequence":"first","affiliation":[{"name":"Ontario Veterinary College, The University of Guelph, Guelph, ON N1G 2W1, Canada"},{"name":"School of Veterinary Science, The University of Queensland, Gatton 4343, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5791-6066","authenticated-orcid":false,"given":"Chiara","family":"Palmieri","sequence":"additional","affiliation":[{"name":"School of Veterinary Science, The University of Queensland, Gatton 4343, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0783-4748","authenticated-orcid":false,"given":"Michael","family":"McGowan","sequence":"additional","affiliation":[{"name":"School of Veterinary Science, The University of Queensland, Gatton 4343, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Frank","family":"Carrick","sequence":"additional","affiliation":[{"name":"School of Environment, The University of Queensland, Gatton 4343, Australia"},{"name":"Sustainable Minerals Institute, Koala Study Program, The University of Queensland, St. Lucia 4072, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-7249-0264","authenticated-orcid":false,"given":"Jackson","family":"Boyd","sequence":"additional","affiliation":[{"name":"School of Veterinary Science, The University of Queensland, Gatton 4343, Australia"},{"name":"School of Environment, The University of Queensland, Gatton 4343, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0290-5458","authenticated-orcid":false,"given":"Stephen D.","family":"Johnston","sequence":"additional","affiliation":[{"name":"School of Veterinary Science, The University of Queensland, Gatton 4343, Australia"},{"name":"School of Environment, The University of Queensland, Gatton 4343, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2023,11,17]]},"reference":[{"key":"ref_1","unstructured":"Forbes, W.A. (1881). Proceedings of the Zoological Society of London, Blackwell Publishing Ltd."},{"key":"ref_2","first-page":"433","article-title":"On the Corpora Lutea and Interstitial Tissue of the Ovary in the Marsupialia","volume":"2","year":"1916","journal-title":"Q. J. Microsc. Sci."},{"key":"ref_3","unstructured":"Mackenzie, W.C. (1919). The Genito-Urinary System in Monotremes and Marsupials, Jenmin, Buxton & Company."},{"key":"ref_4","unstructured":"Handasyde, K.A. (1986). Factors Affecting Reproduction in the Female Koala Phascolarctos cinereus, Monash University."},{"key":"ref_5","unstructured":"Brown, A.S. (1987). Infertility in the Female Koala (Phascolarctos cinereus), University of Queensland."},{"key":"ref_6","unstructured":"Lee, A.K., and Martin, R.W. (1988). The Koala, a Natural History, New South Wales University Press."},{"key":"ref_7","unstructured":"Lee, A.K., and Carrick, F.N. (1989). Fauna of Australia, Vol. 1B, Mammalia, Australian Government Publishing Service."},{"key":"ref_8","unstructured":"Tribe, A., Johnston, S.D., and Carrick, F.N. (1994). Wildlife, Proceedings Number 223, University of Sydney Post Graduate Committee of Veterinary Science."},{"key":"ref_9","first-page":"440","article-title":"Ultrasonographic Diagnosis and Surgical Treatment of Cystic Reproductive Tract Disease in a Female Koala (Phascolarctos cinereus)","volume":"26","author":"Mathews","year":"1995","journal-title":"J. Zoo Wildl. Med."},{"key":"ref_10","unstructured":"Obendorf, D.L. (1988). Australian Wildlife, Proceedings Number 104, University of Sydney."},{"key":"ref_11","unstructured":"Handasyde, K.A., Obendorf, D.L., Martin, R.W., McDonald, I.R., Lee, A.K., and Sanson, G.D. (1990). Biology of the Koala, Surrey Beatty in Association with World Koala Research Corporation. No. March 1991."},{"key":"ref_12","unstructured":"Johnston, S.D. (1999). Studies towards the Development of an Artificial Insemination Protocol in the Koala, Phascolarctos cinereus, The University of Queensland."},{"key":"ref_13","unstructured":"Pagliarani, S. (2021). Further Insights into the Pathogenesis of Chlamydiosis and Characterisation of Immune Cells in the Reproductive Tract of Koalas (Phascolarctos cinereus), The University of Queensland."},{"key":"ref_14","unstructured":"Government, Q. (2022, June 01). Koala Facts. Department of Environment and Science, Available online: https:\/\/environment.des.qld.gov.au\/wildlife\/animals\/living-with\/koalas\/facts."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1530\/jrf.0.1200059","article-title":"Natural and Artificial Methods for Inducing the Luteal Phase in the Koala (Phascolarctos cinereus)","volume":"120","author":"Johnston","year":"2000","journal-title":"J. Reprod. Fertil."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1530\/jrf.0.1200049","article-title":"Studies of the Oestrous Cycle, Oestrus and Pregnancy in the Koala (Phascolarctos cinereus)","volume":"120","author":"Johnston","year":"2000","journal-title":"J. Reprod. Fertil."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1071\/ZO9730001","article-title":"Cyclical Changes in the Uterine Endometrium and Peripheral Plasma Concentrations of Progesterone in the Marsupial Trichosurus Vulpecula","volume":"21","author":"Shorey","year":"1973","journal-title":"Aust. J. Zool."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1002\/ar.1091850306","article-title":"The Growth of the Oocyte and Follicle in the Ovaries of Monotremes and Marsupials","volume":"185","author":"Moore","year":"1976","journal-title":"Anat. Rec."},{"key":"ref_19","unstructured":"Calaby, J.H., and Tyndale-Biscoe, C.H. (1977). Reproduction and Evolution, Australian Academy of Science."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1071\/ZO9850303","article-title":"Effects of Hypophysectomy on Folliculogenesis in the Tammar Macropus eugenii (Marsupialia; Macropodidae)","volume":"33","author":"Panyaniti","year":"1985","journal-title":"Aust. J. Zool."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Tyndale-Biscoe, H., and Renfree, M. (1987). Reproductive Physiology of Marsupials, Cambridge University Press.","DOI":"10.1017\/CBO9780511623493"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1530\/jrf.0.0960755","article-title":"Unexpected Oocyte Growth after Follicular Antrum Formation in Four Marsupial Species","volume":"96","author":"Rodger","year":"1992","journal-title":"J. Reprod. Fertil."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1071\/RD9960661","article-title":"Hormones of Oestrus and Ovulation and Their Manipulation in Marsupials","volume":"8","author":"Hinds","year":"1996","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_24","unstructured":"Parkes, A.S. (1956). The Physiology of Reproduction, Longmans Green and Co."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1530\/jrf.0.0170555","article-title":"Proposal for a Classification of Oocytes and Follicles in the Mouse Ovary","volume":"17","author":"Pedersen","year":"1968","journal-title":"J. Reprod. Fertil."},{"key":"ref_26","unstructured":"Peters, H., and McNatty, K.P. (1980). The Ovary: A Correlation of Structure and Function in Mammals, University of California Press."},{"key":"ref_27","unstructured":"Knobil, E., and Neill, J.D. (1988). The Physiology of Reproduction, Raven Press Ltd."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1530\/rep.1.00300","article-title":"Semen-Induced Luteal Phase and Identification of a LH Surge in the Koala (Phascolarctos cinereus)","volume":"128","author":"Johnston","year":"2004","journal-title":"Reproduction"},{"key":"ref_29","unstructured":"Perry, J.S. (1971). The Ovarian Cycle of Mammals, Oliver and Boyd."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Tyndale-Biscoe, H., and Renfree, M.B. (1987). Reproductive Physiology of Marsupials (Monographs on Marsupial Biology), Cambridge University Press.","DOI":"10.1017\/CBO9780511623493"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1530\/jrf.0.1100361","article-title":"Preovulatory Follicle Development and Ovulation in the Brushtail Possum (Trichosurus vulpecula) Monitored by Repeated Laparoscopy","volume":"110","author":"Crawford","year":"1997","journal-title":"J. Reprod. Fertil."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Austin, C.R. (1961). The Mammalian Egg, Blackwell Publishing.","DOI":"10.5962\/bhl.title.6406"},{"key":"ref_33","unstructured":"Chapman, J. (2003). The Marsupial Zona Pellucida: Its Structure and Glycoconjugate Content. [Ph.D. Thesis, University of Adelaide]."},{"key":"ref_34","first-page":"28","article-title":"Marsupial Gametes and Fertilisation","volume":"5","author":"Rodger","year":"1993","journal-title":"Today\u2019s Life Sci."},{"key":"ref_35","unstructured":"Archer, M. (1987). Possums and Opossums; Studies in Evolution, Royal Zoological Society of New South Wales."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1071\/RD9940451","article-title":"A Comparison of Sperm and Embryo Transport in the Female Reproductive Tract of Marsupial and Eutherian Mammals","volume":"6","author":"Taggart","year":"1994","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_37","unstructured":"Hammond, M.A. (1925). Reproduction in the Rabbit, Oliver and Boyd."},{"key":"ref_38","first-page":"599","article-title":"The Reproductive Cycle of the Water Shrew, Neomys fodiens bicolor (Shaw)","volume":"123","author":"Price","year":"1952","journal-title":"J. Zool."},{"key":"ref_39","unstructured":"Harrison Matthews, L. (1947). Proceedings of the Zoological Society of London, Blackwell Publishing Ltd."},{"key":"ref_40","unstructured":"Woolley, P. (1966). Reproductive Physiology of Antechinus Stuartii Macleay. [Ph.D. Thesis, Australian National University]."},{"key":"ref_41","first-page":"17","article-title":"Observations on the Histology of the Opossum Ovary","volume":"30","year":"1942","journal-title":"Contrib. Embryol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1457","DOI":"10.1071\/RD19034","article-title":"Ovary of the Southern Hairy-Nosed Wombat (Lasiorhinus latifrons): Its Divergent Structural Organisation","volume":"31","author":"Breed","year":"2019","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"R157","DOI":"10.1530\/REP-13-0012","article-title":"Wombat Reproduction (Marsupialia; Vombatidae): An Update and Future Directions for the Development of Artificial Breeding Technology","volume":"145","author":"Hogan","year":"2013","journal-title":"Reproduction"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1002\/aja.1000610205","article-title":"The Thecal Gland and Its Relation to the Reproductive Cycle. A Study of the Cyclic Changes in the Ovary of the Pocket Gopher, Geomys bursarius (Shaw)","volume":"61","author":"Mossman","year":"1937","journal-title":"Am. J. Anat."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1002\/ar.1090830202","article-title":"The Thecal Gland in the Guinea Pig Ovary","volume":"83","author":"Stafford","year":"1942","journal-title":"Anat. Rec."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Austin, C.R., and Short, R.V. (1984). Reproduction in Mammals, Book 3: Hormonal Control of Reproduction, Cambridge University Press.","DOI":"10.1017\/CBO9781139167727"},{"key":"ref_47","unstructured":"Lamming, G.E. (1984). Marshall\u2019s Physiology of Reproduction, Churchill Livingstone."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1210\/endo-82-1-122","article-title":"Effect of Prolactin on Progestin Release and Cholesterol Storage by Rabbit Ovarian Interstitium","volume":"82","author":"Hilliard","year":"1968","journal-title":"Endocrinology"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"901","DOI":"10.1210\/endo-80-5-901","article-title":"A Functional Role for 20-Alpha-Hydroxypregn-4-En-3-One in the Rabbit","volume":"80","author":"Hilliard","year":"1967","journal-title":"Endocrinology"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"852","DOI":"10.1210\/endo-99-3-852","article-title":"Ovarian Regulation of Postcoital Gonadotropin Release in the Rabbit: Reexamination of a Functional Role for 20 \u03b1Dihydroprogesterone","volume":"99","author":"Goodman","year":"1976","journal-title":"Endocrinology"},{"key":"ref_51","unstructured":"Lee, A.K., Handasyde, K.A., and Sanson, G. (1990). Biology of the Koala, Surrey Beatty & Sons Pty Ltd."},{"key":"ref_52","unstructured":"Knobil, E., and Neill, J.D. (1994). The Physiology of Reproduction, Raven Press Ltd."},{"key":"ref_53","unstructured":"Hafez, E.S.E., and Bladau, R.J. (1969). The Mammalian Oviduct, University of Chicago Press."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1071\/ZO9720235","article-title":"Uterine Glandular Regeneration during the Follicular Phase in the Marsupial Trichosurus vulpecula","volume":"20","author":"Shorey","year":"1972","journal-title":"Aust. J. Zool."},{"key":"ref_55","first-page":"513","article-title":"The Female Urogenital System of Trichosurus vulpecula (Marsupialia)","volume":"24","author":"Kean","year":"1964","journal-title":"Aust. J. Zool."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1071\/ZO9630250","article-title":"Distribution and Comparative Histology of Cutaneous Glands in Certain Marsupials","volume":"11","author":"Green","year":"1963","journal-title":"Aust. J. Zool."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1071\/AM84018","article-title":"Observations on the Reproductive Systems and Paracloacal Glands of Cercartetus lepidus (Marsupialia: Burramyidae)","volume":"7","author":"Smith","year":"1984","journal-title":"Aust. Mammol."},{"key":"ref_58","unstructured":"Smith, A.P., and Hume, I.D. (1984). Possums and Gliders, Australian Mammal Society."},{"key":"ref_59","unstructured":"Fawcett, D., and Raviola, E. (1986). Bloom and Fawcett, a Textbook of Histology, Chapman and Hall. [12th ed.]."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1002\/jez.1400940307","article-title":"Embryonic Differentiation of Cowper\u2019s and Bartholin\u2019s Glands of the Opossum Following Castration and Ovariotomy","volume":"94","author":"Rubin","year":"1943","journal-title":"J. Exp. Zool."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1002\/jmor.1050740202","article-title":"The Relation of Hormones to the Development of Cowper\u2019s and Bartholins Glands in Opossum (Didelphys Virginiana)","volume":"74","author":"Rubin","year":"1944","journal-title":"J. Morphol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1002\/ar.1091010206","article-title":"A Study of the Bovine Vestibular Gland","volume":"101","author":"Reutner","year":"1948","journal-title":"Anat. Rec."},{"key":"ref_63","unstructured":"Lee, A.K., Handasyde, K.A., and Sanson, G.D. (1990). Biology of the Koala, Surrey Beatty and Sons Pty Ltd."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"426","DOI":"10.1053\/j.semperi.2015.07.001","article-title":"Overview of the Mechanisms of Induction of Labor","volume":"39","author":"Irani","year":"2015","journal-title":"Semin. Perinatol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"160","DOI":"10.1111\/j.1748-1090.2003.tb02076.x","article-title":"Birth of Koalas (Phascolarctos cinereus) at Lone Pine Koala Sanctuary Following Artificial Insemination","volume":"38","author":"Johnston","year":"2003","journal-title":"Int. Zoo Yearb."}],"container-title":["Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2079-7737\/12\/11\/1445\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T21:24:47Z","timestamp":1760131487000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2079-7737\/12\/11\/1445"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,11,17]]},"references-count":65,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2023,11]]}},"alternative-id":["biology12111445"],"URL":"https:\/\/doi.org\/10.3390\/biology12111445","relation":{},"ISSN":["2079-7737"],"issn-type":[{"value":"2079-7737","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,11,17]]}}}