## 4 Discussion  
### Horizontal Movements, Habitat Use, Revisitations, and Home Ranges  
The reef manta rays tracked via satellite in our study exhibited localized movements within and around the respective tagging areas in Indonesian waters. In Komodo, the localized movements of the satellite-tracked individuals were consistent with the findings from a passive acoustic telemetry study of the same species conducted over a decade earlier [10]. In the Raja Ampat region, the movement patterns reported in the current study closely resemble those observed using passive acoustic telemetry on the same species, where frequent movements occurred between aggregation sites 5-12 km apart, with occasional long-distance movements [12]. Comparable localized movement patterns have also been documented for the reef manta rays tracked via satellite in other regions, including the  
Figure 4: Home range (95% utilization distribution) of satellite-tracked reef manta rays in Berau, northern and southern Raja Ampat, Nusa Penida, and Komodo from July 2014 to July 2022. Red Sea [18], Dungonab Bay and Mukkawar Island National Park, Sudan [45], eastern Australia [16], western Australia [46], and New Caledonia [19]. While the satellite telemetry data in our study align with the existing knowledge from passive acoustic tracking [10; 12], our approach offers unique contributions. Satellite telemetry provides spatial and temporal resolution that complements passive acoustic tracking by covering broader geographic areas and capturing behaviors beyond the range of acoustic receiver arrays [13]. Additionally, it helps identify key sites that may not have been previously identified, expanding our understanding of habitat use and movement patterns. These contributions underscore the value of our study in advancing reef manta ray ecology. ARS behaviors were observed in a number of areas in our study regions in Indonesia (Figure 2). The areas in each region where ARS behaviors were observed and revisited by the satellite-tracked reef manta rays are known as reef manta ray aggregation sites, frequently used for either cleaning, feeding, or both. The reefs around Sangalaki Island in Berau have both cleaning sites and feeding sites for the reef manta population in the region [25], and Sangalaki is the only area identified in Berau and the broader East Kalimantan region with frequent sightings of reef manta rays [25]. The relatively long average visitation time (34.8 h) and high revisitations at Sangalaki highlight the ecological importance of this site for the vulnerable species. The southwest coast of Nusa Penida, including Manta Point, has long been known as an important area for feeding and cleaning by reef manta rays [8], and our satellite tagging results validate this significance. Similarly, Karang Makassar in Komodo has been previously highlighted as an important feeding and cleaning area for reef manta rays based on photo ID [7] and acoustic telemetry [10], again confirmed by our results herein. Importantly, Baru Bay on the east of Rinca Island has not previously been identified as an important area for reef manta rays based on those two previous manta ray studies in the region [7; 10], and our findings here suggest that additional survey efforts should be expended in eastern Rinca to further investigate the use of this area by reef manta rays. Finally, the areas highlighted as important aggregation sites for reef manta rays in the current study in Raja Ampat largely confirm the previous findings reported from studies using long-term photo ID and passive acoustic telemetry [6; 11; 12]. Nonetheless, the recursive analysis of Raja Ampat satellite-tracking data did highlight several new areas to the east of Dayan Island in northern Batanta (Figure 3) that have not been documented previously as important areas for manta rays, and further investigation is needed to confirm these observations. The current study also further supports the importance of Ayau Besar Lagoon in northern Raja Ampat as an important area for reef manta rays [6; 12].