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Background: Biodiversity is being lost rapidly and its conservation is thus one of the most urgent tasks today. For biodiversity conservation to be successful, the public needs to gain an awareness and understanding of biodiversity and its importance. Moreover, species experts are needed who have the skills necessary for identifying and recording biodiversity. Previous research showed that citizen science projects can contribute to educating the public about biodiversity. However, it is still unclear how project characteristics connect to participants' knowledge and skills and how citizen science projects should be designed if they are to foster participants' learning. Aim: We aimed to investigate specific characteristics of biodiversity citizen science projects that could potentially influence participants' learning. We explored the following project characteristics from both the project coordinators' and the participants' perspectives: information and training provided to participants, social interaction among participants, contact between participants and staff, and feedback and recognition provided to participants. Methods and results: In order to examine the extent to which these project characteristics are connected to participants' gains in knowledge and skills, we conducted a comprehensive study across 48 biodiversity citizen science projects in Europe and Australia. We found that participants' perceived gains in knowledge and skills were significantly related to the five project characteristics as reported by the participants: information received by the participants, training received by the participants, social interaction among participants, contact between participants and staff, and feedback and recognition received by the participants. Conclusion: We conclude that by deliberately designing citizen science projects to include features such as interaction and feedback, these projects could achieve higher learning outcomes for the participants. Thereby, suitable modes of communication between ...