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Abstract Scleractinian corals provide habitats for a broad variety of cryptofauna, which in turn may contribute to the overall functioning of coral symbiomes. Among these invertebrates, hydrozoans belonging to the genus Zanclea represent an increasingly known and ecologically important group of coral symbionts. In this study, we analysed 321 Zanclea colonies associated with 31 coral genera collected from 11 localities across the Indo-Pacific and Caribbean regions, and used a multi-disciplinary approach to shed light on the evolution and biogeography of the group. Overall, we found high genetic diversity of hydrozoans that spans nine clades corresponding to cryptic or pseudo-cryptic species. All but two clades are associated with one or two coral genera belonging to the Complex clade, whereas the remaining ones are generalists associated with both Complex and Robust corals. Despite the observed specificity patterns, no congruence between Zanclea and coral phylogenies was observed, suggesting a lack of coevolutionary events. Most Zanclea clades have a wide distribution across the Indo-Pacific, including a generalist group extending also into the Caribbean, while two host-specific clades are possibly found exclusively in the Red Sea, confirming the importance of this peripheral region as an endemicity hotspot. Ancestral state reconstruction suggests that the most recent common ancestor of all extant coral-associated Zanclea was a specialist species with a perisarc, occurring in what is now known as the Indo-Pacific. Ultimately, a mixture of geography- and host-related diversification processes is likely responsible for the observed enigmatic phylogenetic structure of coral-associated Zanclea. ; The authors wish to thank all the people involved in collecting material or organising sampling campaigns: Tullia Isotta Terraneo (KAUST, Saudi Arabia), Malek Amr Gusti (KAUST, Saudi Arabia), Timothy Ravasi (OIST, Japan), the captain and crew of the MV Dream-Master (Saudi Arabia), the KAUST Coastal and Marine Resources Core Lab, Inga Dehnert (UNIMIB, Italy), Nicholas WL Yap (NUS, Singapore), Sudhanshi S Jain (NUS, Singapore), Stephen Keable (Australian Museum), Penny Berents (Australian Museum), Anne Hoggett (Australian Museum), Lyle Vail (Australian Museum), Masayuki Tamae (Marine Space, Japan), Hiroki Nakakouji (Marine Space, Japan), Yoshihiro Katsushima (Win Diving, Japan), Yuting V Lin (IONTU, Taiwan), Jian-Wen Chen (IONTU, Taiwan), and staff at the Green Island Marine Research Station (Taiwan) and Academia Sinica (Taiwan). Permissions relevant to undertake the research have been obtained from the applicable governmental agencies. The work was partially funded by PADI Foundation Grants #28634 and #14384 to DM and SM, and baseline research funds of PG and MLB. Samples from Eilat (Israel) were collected during the HyDRa Project funded by the EU FP7 Research Infrastructure Initiative 'ASSEMBLE' (Grant #227799) to DP. Samples from Ludao (Taiwan) were collected under the permit #1060174405 issued by Taitung County Government with the support of a grant from the Ministry of Science and Technology of Taiwan (#106-2611-M-002-008) to VD. Financial support to DP for collecting samples at Lizard Island (Australia) was provided by the 2018 John and Laurine Proud Fellowship and the Australian Museum's Lizard Island Research Station. Fieldwork in Singapore was partially funded by the National Research Foundation, Prime Minister's Office, Singapore under its Marine Science R&D Programme (MSRDP-P03) to DH.

11 pages, 3 figures, 1 table, supplementary material https://doi.org/10.1126/sciadv.abm7452.-- Data and materials availability: All data needed to evaluate the conclusions in the paper are present in the paper, the Supplementary Materials, Dryad, and Zenodo ; Phylogenetic relationships and the timing of evolutionary events are essential for understanding evolution on longer time scales. Cheilostome bryozoans are a group of ubiquitous, species-rich, marine colonial organisms with an excellent fossil record but lack phylogenetic relationships inferred from molecular data. We present genome-skimmed data for 395 cheilostomes and combine these with 315 published sequences to infer relationships and the timing of key events among c. 500 cheilostome species. We find that named cheilostome genera and species are phylogenetically coherent, rendering fossil or contemporary specimens readily delimited using only skeletal morphology. Our phylogeny shows that parental care in the form of brooding evolved several times independently but was never lost in cheilostomes. Our fossil calibration, robust to varied assumptions, indicates that the cheilostome lineage and parental care therein could have Paleozoic origins, much older than the first known fossil record of cheilostomes in the Late Jurassic ; This project is funded by the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant agreement no. 724324 to L.H.L.). Extra funding for specific sampling expeditions were provided by the National Science Centre of Poland (grant PANIC/2016/23/B/ST10/01936 to P.K.), the ASSEMBLE Plus (Horizon 2020), the Leverhulme Trust (Research Project Award RPG-2016-429 to A.W.), the Russian Science Foundation (grant 18-14-00086 to A.N.O.), Conselho Nacional de Desenvolvimento Científico e Tecnológico, Brazil [PQ-CNPq 308768/2018-3], Fundação de Amparo à Pesquisa do Estado de São Paulo, Brazil [FAPESP 19/17721-9] and the DISTANTCOM project (CTM2013-42667/ANT to C. Avila). B.F. was supported by Beatriu de Pinós (2019-BP-00183), funded by the AGAUR (Government of Catalonia) and by the Horizon 2020 programme under the Marie Skłodowska-Curie grant (no. 801370). The CAML-CEAMARC cruise of RV Aurora Australis (IPY project no. 53) were supported by the Australian Antarctic Division, the Japanese Science Foundation, the French Polar Institute IPEV, and the Muséum national d'Histoire naturelle, led by voyage leader, M. Riddle ; With the institutional support of the 'Severo Ochoa Centre of Excellence' accreditation (CEX2019-000928-S) ; Peer reviewed