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While some of these components have been lost during evolution, others remain undetectable by standard bioinformatics approaches. Using a conditional splitCas9 system in Toxoplasma gondii, we previously identified TGGT1_301410, a hypothetical gene conserved among apicomplexans, as a potential trafficking factor. Here, we show that TGGT1_301410 is a distant ortholog of T. gondii tepsin (TgTEP), localized to the trans-Golgi and functioning as an accessory protein of the adaptor protein complex 4 (AP4). We demonstrate that AP4-TgTEP is essential for the actin-dependent transport of vesicles to the plant-like vacuole (PLVAC) and Golgi organization. Notably, our findings reveal that, unlike in metazoans, the AP4 complex in T. gondii utilizes clathrin as a coat protein, a mechanism more closely aligned with that of plants. These results underscore a conserved yet functionally adapted vesicular transport system in Apicomplexa.<\/jats:p>","DOI":"10.1083\/jcb.202312109","type":"journal-article","created":{"date-parts":[[2025,6,13]],"date-time":"2025-06-13T02:21:04Z","timestamp":1749781264000},"update-policy":"https:\/\/doi.org\/10.1083\/jcb.crossmarkpolicy","source":"Crossref","is-referenced-by-count":1,"title":["Tepsin and AP4 mediate transport from the trans-Golgi to the plant-like vacuole in toxoplasma"],"prefix":"10.1083","volume":"224","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8641-3313","authenticated-orcid":true,"given":"Janessa","family":"Grech","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05591te55","id-type":"ROR","asserted-by":"publisher"}],"name":"Ludwig-Maximilians-Universit\u00e4t, LMU 1 Experimental Parasitology, Department of Veterinary Sciences, Faculty of Veterinary Medicine, , 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